LELAND   STANFORD   JUNIOR  UNIVERSITY  PUBLICATIONS 
UNIVERSITY  SERIES 


DUDLEY  MEMORIAL 
VOLUME 


CONTAINING  A  PAPER 

BY 

WILLIAM  RUSSEL  DUDLEY 

AND 

APPRECIATIONS  AND  CONTRIBUTIONS 

IN  HIS  MEMORY 

BY 

ERIENDS  AND  COLLEAGUES 


(WITH  PORTRAIT) 


STANFORD  UNIVERSITY,  CALIFORNIA 

PUBLISHED  BY  THE  UNIVERSITY 

1913 


crfftjus 


BIO-AGRICULTURAL  LIBRARY 
UNIVERSITY  OF  CALIFORNIA 
RIVERSIDE,  CALIFORNIA  92521 


WILLIAM     RUSSEL     DUDLEY 


LELAND  STANFORD    JUNIOR  UNIVERSITY  PUBLICATIONS 
UNIVERSITY  SERIES 


DUDLEY  MEMORIAL 
VOLUME 


CONTAINING  A  PAPER 

BY 

WILLIAM  RUSSEL  DUDLEY 

AND 

APPRECIATIONS  AND  CONTRIBUTIONS 

IN  HIS  MEMORY 

BY 

FRIENDS  AND  COLLEAGUES 


(WITH  PORTRAIT) 

Library 

Experiment  Siaticc. 
of  California 


STANFORD  UNIVERSITY,  CALIFORNIA 

PUBLISHED  BY  THE  UNIVERSITY 

1913 


•     /  7/3 


TABLE  OF  CONTENTS 
WILLIAM  RUSSEL  DUDLEY 5 

MEMORIAL  ADDRESSES: 

JOHN  CASPER  BRANNER 7 

DOUGLAS  HOUGHTON  CAMPBELL 1 1 

APPRECIATIONS  : 

DAVID  STARR  JORDAN 16 

LER.OY  ABRAMS  20 

GEORGE  JAMES  PEIRCE 22 

JARED  TREMAN  NEWMAN 23 

WILLIAM  FRANKLIN  WIGHT 25 

LIST  OF  PUBLICATIONS  OF  W.  R.  DUDLEY 27 

LIST  OF  CORNELL  UNIVERSITY  PUPILS  OF  W.  R.  DUDLEY 29 

LIST  OF  STANFORD  UNIVERSITY  PUPILS  OF  W.  R.  DUDLEY 30 

SCIENTIFIC  PAPERS: 

THE  VITALITY  OF  SEQUOIA  GIGANTEA 33 

WILLIAM  RUSSEL  DUDLEY,  LATE  PROFESSOR  OF  BOTANY 

THE  MORPHOLOGY  AND  SYSTEMATIC  POSITION  OF  CALYCULARIA 

RADICULOSA    (STEPH.)    (TWELVE   FIGURES)    43 

DOUGLAS  HOUGHTON  CAMPBELL,  PROFESSOR  OF  BOTANY 

STUDIES   OF   IRRITABILITY   IN   PLANTS,  III,  THE   FORMATIVE 

INFLUENCE  OF  LIGHT  (ONE  PLATE) 62 

GEORGE  JAMES  PEIRCE,  PROFESSOR  OF  BOTANY  AND  PLANT 
PHYSIOLOGY 

THE  GYMNOSPERMS  GROWING  ON  THE  GROUNDS  OF  STANFORD 

UNIVERSITY  (Six  PLATES) 81 

LEROY  ABRAMS,  ASSOCIATE  PROFESSOR  OF  BOTANY 

THE  SYNCHYTRIA  IN  THE  VICINITY  OF  STANFORD  UNIVERSITY 

(ONE  PLATE) in 

JAMES  McMuRPHY,  INSTRUCTOR  IN  SYSTEMATIC  BOTANY 

THE  LAW  OF  GEMINATE  SPECIES 115 

DAVID  STARR  JORDAN,  PRESIDENT  OF  STANFORD  UNIVERSITY 

SOME  RELATIONS  BETWEEN  SALT  PLANTS  AND  SALT-SPOTS.  .  .     123 
WILLIAM  AUSTIN  CANNON,  DESERT  LABORATORY 

NORTH  AMERICAN  SPECIES  OF  THE  GENUS  AMYGDALUS 130 

WILLIAM  FRANKLIN  WIGHT,  BUREAU  OF  PLANT  INDUSTRY 


WILLIAM  RUSSEL  DUDLEY  was  born  at  Guil- 
ford,  Connecticut,  March  1,  1849,  and  died  at 
Los  Altos,  California,  June  4,  1911.  He  was 
educated  at  Cornell  University,  graduating  with  the  class  of 
1874,  and  taking  his  master's  degree  at  the  same  institution 
in  1876.  He  was  instructor  in  botany  in  Cornell  Univer- 
sity from  1873  to  1876,  and  assistant  professor  of  botany 
from  1876  to  1892.  On  leave  of  absence  from  Cornell  he 
was  acting  professor  of  biology  in  the  University  of  Indiana 
in  1880,  and  he  spent  the  year  1886-1887  studying  at  Stras- 
burg  and  Berlin.  He  was  appointed  professor  of  system- 
atic botany  at  Stanford  University  in  1893,  a  position  which 
he  held  from  that  time  until  December,  1910,  when,  on  ac- 
count of  ill  health,  he  voluntarily  retired  and  became  pro- 
fessor emeritus. 


Unfoersfttp  Cfjapel 

Septemfter  10,  1911 


Jfflemortam 

MtUtam  l\ussd 

(Emeritus  professor  of  IBotanp 


Jiorn  jWarcft  I,  1849       BtebSTune  4,  1911 


WILLIAM  RUSSEL  DUDLEY 

[An  address  delivered  at  the  services  held  in  the  University  Chapel  of 
Stanford  University,  September  10,  1911.] 

By  JOHN  CASPER  BRANNER,  Vice- President 

I  DOUBT  if  there  is  any  time  in  men's  lives  when  they  come  to  know  each 
other  as  well  as  they  do  in  their  college  days,  especially  when  they 
happen  to  have  the  same  studies,  to  be  in  the  same  classes,  and  to 
be  much  thrown  together  by  any  circumstances  whatever. 

Professor  Dudley  and  I  belonged  to  the  class  of  1874  in  Cornell,  we 
had  some  of  the  same  studies,  we  belonged  to  the  same  fraternity,  and  as 
students  we  had  about  the  same  ups  and  downs.  Aside  from  these  mutual 
interests  we  were  thrown  together  still  more  by  the  fact  that  Dudley, 
being  a  student  in  the  scientific  course,  had  botany  in  the  early  part  of  his 
studies,  while  I  in  the  course  in  Greek  and  Latin  took  botany  near  the 
end  of  my  college  work,  and  so  it  came  about  that  in  our  senior  year  he 
was  instructor  in  botany  and  I  was  his  pupil. 

As  enthusiastic  students  and  as  intimate  friends  we  tramped  together 
every  hill,  explored  every  gorge  and  penetrated  every  swamp  for  many 
miles  around  Ithaca.  Under  his  guidance  I  came  to  have  a  personal 
acquaintance  with  and  affection  for  every  flowering  plant  of  the  region 
about  Cayuga  Lake,  and  for  Dudley  always  a  deeper  love  and  a  greater 
esteem. 

The  first  piece  of  scientific  work  I  ever  did — a  study  of  the  fibro- 
vascular  bundles  in  the  palms — was  undertaken  and  carried  through  under 
his  guidance. 

On  the  slopes  of  the  hills  west  of  Ithaca  it  was  he  who  pointed  out 
to  me  for  the  first  time  the  deep  marks  cut  in  the  hard  rocks  by  the  ice 
of  the  glacial  epoch.  Thus  Dudley  was  not  only  my  first  and  principal 
instructor  in  botany,  but  he  was  also,  in  a  way,  my  first  effective  instructor 
in  geology. 

We  college  professors  are  more  or  less  given  to  the  discussion  of  methods 
of  instruction,  and  it  is  no  uncommon  thing  to  hear  this  or  that  man's 
methods  found  fault  with.  I  dare  say  such  criticisms  are  well  enough  in 
their  way,  but  after  all  is  said  and  done  there  remains  one  supreme  test 
of  a  teacher  that  is  often  lost  sight  of  in  these  discussions,  and  that  is 
his  results.  I  do  not  speak  with  a  knowledge  of  the  precise  number  of 
his  students  who  stand  to-day  in  the  front  rank  of  our  botanists,  but  my 
general  impression  is  that,  judged  by  this  standard — by  results  with  his 


8  DUDLEY    MEMORIAL   VOLUME 

students — Professor  Dudley  was  one  of  the  most  successful  teachers  of 
botany  this  country  has  ever  produced.  And  I  am  confident  that  that 
success  is  to  be  attributed  to  a  great  extent  to  the  human  and  personal 
rather  than  to  the  technical  part  of  his  methods  assa  teacher. 

He  wa£  always  at  the  service  of  his  students.  No  hour  of  the  day 
or  the  night  was  inopportune  when  a  student  wanted  his  advice  or  direction. 

His  personal  influence  during  his  early  manhood  was  the  finest  and 
most  wholesome  that  I  have  ever  found  among  men,  whether  old  or  young. 
Professor  W.  R.  Lazenby  of  the  University  of  Ohio,  who  was  a  classmate, 
writes  of  him:  "I  may  say  for  myself  that  I  owe  Dudley  a  great  deal. 
I  roomed  with  him  my  first  year  at  Cornell,  and  he  had  a  great  influence 
for  good  over  my  life.  I  think,  all  in  all,  he  was  one  of  the  best  men 
I  ever  knew — pure  gold." 

Dudley  was  a  warm  hearted,  genuine  lover  of  nature  in  all  her  forms 
and  in  all  her  moods,  and  this  gave  him  that  enthusiasm  without  which  a 
teacher  is  not  a  teacher.  No  man  could  have  fitted  more  perfectly  into 
the  sentimental  side  of  botany — if  botany  has  any  such  side.  The  colors, 
the  beauty,  and  delicacy  of  flowers  and  plants,  their  lives,  their  kinships, 
their  histories — all  appealed  to  the  artistic  side  of  his  nature. 

This  love  for  and  appreciation  of  nature,  however,  was  his  despair 
as  well  as  his  constant  delight.  His  soul  overflowed  with  affection  for  it 
all,  but  he  was  so  sensitive  to  the  defects  of  language  and  of  other  methods 
of  representation  that  he  rarely  undertook  to  give  expression  to  his  love 
for  it. 

But  I  would  not  have  you  imagine  that  he  was  a  botanist  and  nothing 
but  a  botanist,  neither  was  he  a  scientific  man  to  the  exclusion  of  other 
interests.  Indeed  he  was  deeply  and  generally  interested  in  everything 
human  and  spiritual. 

At  heart  he  was  a  poet.  I  shall  never  forget  the  glow  of  enthusiasm 
with  which  he  read  to  me,  when  it  first  appeared,  Longfellow's  Morituri 
Salutamus.  He  always  had  about  him  the  works  of  the  best  poets  and  a 
few  pictures  and  other  works  of  art  of  the  first  quality. 

His  was 

"The  love  of  learning,  the  sequestered  nooks, 
And  all  the  sweet  serenity  of  books." 

To  be  rather  than  to  appear  was  the  steadfast  principle  of  his  life. 
Modesty,  gentleness,  unobtrusiveness,  decorum,  and  purity  of  life  were  his 
most  prominent  characteristics.  He  never  did  anything  for  the  sake  of 
display;  he  never  courted  popularity.  His  whole  life,  within  and  without, 
was  one  long,  living  protest  against  vulgarity  in  all  its  forms. 


WILLIAM    RUSSEL    DUDLEY BRANNER 

He  was  a  man  of  the  finest  possible  fiber,  so  fine  indeed  that  the  very 
delicacy  of  his  nature  unfitted  him  for  some  of  the  pioneer  work  he  was 
called  upon  to  do  in  his  lifetime. 

When  Dr.  Jordan  was  President  of  the  University  of  Indiana,  he  tried 
for  some  time  to  induce  Dudley  to  go  to  that  institution  as  professor  of 
botany.  And  I  recall  in  this  connection  that  Dr.  Jordan  said  to  me  on 
one  occasion:  "Quite  aside  from  his  ability  as  a  teacher  of  botany  we  need 
him  here  on  account  of  his  personal  influence." 

But  Dudley  declined  the  proffered  position  largely  because  he  felt 
that  he  was  not  altogether  fitted  for  the  pioneer  work  required  there  at 
that  time. 

With  the  idea  that  poverty  helps  rather  than  hinders  a  young  man, 
Dudley  did  not  altogether  agree;  in  fact  he  entirely  disagreed  with  it  in 
so  far  as  it  related  to  himself.  He  felt  keenly  the  inconveniences  of  having 
to  earn  his  living  while  carrying  on  his  studies. 

The  necessity  of  devoting  so  much  time  to  his  teaching  and  the 
strictness  of  the  standards  he  set  for  himself  explain  why  he  was  not  a  writer 
of  books  or  the  publisher  of  a  very  long  list  of  scientific  papers. 

Lest  some  who  did  not  know  him  well  should  imagine  that  so  much 
self-effacement  indicated  a  man  with  but  little  force  of  character  I  hasten 
to  say  that  such  was  very  far  from  being  the  case.  With  all  his  gentleness 
and  sweetness  I  have  never  known  a  man  of  more  decision  of  character, 
stronger  will  power,  or  of  more  determination,  firmness,  and  unswerving 
purpose. 

In  the  summer  of  1882,  I  think  it  was,  when  Dudley  was  thirty-three 
years  old,  the  baccalaureate  sermon  at  Cornell  was  preached  by  the  Rev. 
Dr.  Heber  Newton,  who  was  for  a  while  chaplain  here  at  Stanford  Uni- 
versity. 

Dudley  and  I  went  to  chapel  together.  We  found  it  so  crowded  that 
we  could  not  find  seats  together,  and  I  sat  in  the  row  of  seats  just  behind 
him.  Dr.  Newton's  address  was  a  eulogy  upon  the  life,  character,  and 
influence  of  Ralph  Waldo  Emerson.  You  can  imagine  the  tribute  he  paid 
to  that  distinguished  writer  and  lecturer.  I  recall  that  when  Dr.  Newton 
had  finished  his  eloquent  address  I  said  to  myself:  "Yes,  but  right  here 
living  in  our  own  midst  and  within  the  reach  of  my  hand  is  a  man  who 
has  every  one  of  the  finest  traits  of  character  of  Emerson." 

In  the  latter  part  of  his  life  certain  of  his  traits  became  more  prominent 
than  during  his  younger  manhood.  He  was  always,  and  of  necessity,  a 
purist  in  every  sense  in  which  that  word  can  be  used.  But  as  he  grew 
older  I  imagine  that  his  sensitiveness  brought  him  more  pain  than  pleasure, 


10  DUDLEY    MEMORIAL   VOLUME 

and  to  this  I  attribute  the  rather  lonely  life  he  led  after  coming  to 
California. 

Unfortunately  there  are  those  who  knew  Professor  Dudley  only  as  a 
name  in  the  university  register.  I  am  sure  my  friend  would  not  thank 
me  to  apologize  for  the  modest  part  he  played  in  this  or  in  any  other 
community,  but  in  closing  I  am  constrained  to  say  a  word  in  behalf  of 
him  and  of  all  such  men:  It  behooves  us  not  to  lose  sight  of  this  blessed 
truth,  that  there  are  fine  men  and  women  in  this  world  of  ours — and  plenty 
of  them,  too — who  keep  out  of  the  limelights,  whose  names  we  never  see 
in  the  headlines  of  the  newspapers,  but  who  lead  quiet,  sane,  and  wholesome 
lives.  Such  people  always  suggest  to  me  the  foundations  of  a  great  structure. 
These  foundations  lie  deep  beneath  the  surface  of  the  ground;  we  never 
see  them;  we  seldom  think  of  them;  they  are  not  decorated  with  flying 
flags  or  written  across  with  gaudy  colors  or  blazing  electric  lights..  But 
they  stand  fast  and  firm,  and  the  stability  and  the  real  worth  of  the  entire 
superstructure  depends  upon  them. 

One  of  these  foundation-men  was  William  Russel  Dudley. 


WILLIAM  RUSSEL  DUDLEY 

[An  address  delivered  at  the  services  held  in  the  University  Chapel  of 
Stanford  University,  September  10,  1911.] 

By  PROFESSOR  DOUGLAS  HOUGHTON  CAMPBELL 

WILLIAM  RUSSEL  DUDLEY  was  born  in  Guilford,  Connecticut,  in  1849 
and  was  one  of  the  earlier  students  of  Cornell  University,  from 
which  he  graduated  with  the  degree  of  Bachelor  of  Science  in 
1874.  At  that  time  Cornell  University  had  only  been  opened  for  a  short 
time,  and  I  fancy  the  conditions  there  were  in  many  respects  very  much 
like"  those  of  Stanford  twenty  years  ago.  The  new  university  at  Ithaca 
had  broken  away  from  the  traditions  of  the  earlier  eastern  colleges,  and 
science  received  far  more  attention  than  in  most  of  the  other  institutions. 
The  opening  of  the  new  university  with  its  facilities  for  scientific  work 
attracted  a  group  of  young  men  who  have  since  attained  pre-eminence  in 
their  various  departments'.  Among  those  who  are  on  our  own  faculty  were 
Dr.  Jordan  and  Professor  Branner,  with  whom  Professor  Dudley  was  asso- 
ciated on  intimate  terms.  Of  Professor  Dudley's  life  as  spent  at  Cornell, 
Professor  Branner  has  just  given  us  a  most  sympathetic  account.  In  Dr. 
Jordan's  recent  sketch  of  Professor  Dudley  in  Science,  he  tells  us  that  for  a 
time  he  was  himself  instructor  in  botany,  and  that  Professor  Dudley  during 
the  early  part  of  his  stay  at  Cornell  came  under  his  instruction.  However, 
it  was  not  long  before  Dudley  himself  was  acting  as  instructor  even  in  his 
undergraduate  days,  and  later  became  attached  to  the  staff  of  the  university. 
It  is  hard  for  us  to  realize  in  these  days  when  every  college  or 
university  of  any  pretensions  whatever  has  its  department  of  botany  well- 
equipped  and  well-manned,  that  during  the  '70s  the  number  of  profes- 
sorships of  botany  in  the  whole  United  States  probably  did  not  exceed 
half  a  dozen.  Cornell  was  one  of  the  first  of  the  universities  to  establish 
a  distinct  chair  of  botany,  and  at  the  time  that  Professor  Dudley  entered 
Cornell  the  chair  was  held  by  Professor  Albert  Prentiss.  While  a  student 
at  Cornell,  Dudley  attended  the  summer  session  of  the  famous  school  at 
Penikese  where  Agassiz  for  the  first  time  instituted  a  seaside  summer 
school,  the  model  of  which  has  since  been  repeated  in  so  many  places.  At 
Penikese  Dudley  was  associated  not  only  with  his  fellow  students  of  Cornell 
but  also  with  a  number  of  other  men  who  laid  the  foundation  of  the  biological 
studies  which  have  had  such  a  tremendous  influence  in  the  development  of 
science  since  that  time. 


12  DUDLEY    MEMORIAL    VOLUME 

Professor  Dudley  very  early  became  interested  in  the  problem  of  plant 
distribution.  The  region  about  Ithaca  is  a  peculiarly  interesting  one  botani- 
cally,  offering  an  unusual  variety  of  conditions  with  a  correspondingly 
varied  and  interesting  flora.  Dudley  soon  became  intimately  acquainted 
with  the  flora  of  this  whole  region  and  the  results  of  his  studies  were 
later  published  under  the  name  "The  Cayuga  Flora."  This  was  soon  sup- 
plemented by  a  second  similar  work  on  "The  Lackawanna  and  Wyoming 
Flora."  While  at  Cornell,  Professor  Dudley  also  published  in  collaboration 
with  Professor  M.  B.  Thomas  a  "Manual  of  Histology."  He  also  published 
a  number  of  other  shorter  papers  dealing  mainly  with  the  flora  of  the 
same  region. 

During  the  latter  part  of  his  stay  at  Cornell  Professor  Dudley  had 
charge  of  the  work  on  the  lower  plants,  especially  the  fungi,  to  which  he 
devoted  much  attention.  In  connection  with  this  work  upon  the  fungi 
Professor  Dudley  made  a  trip  to  Europe  in  1887,  and  it  was  upon  this 
trip  that  I  had  the  first  opportunity  of  making  his  acquaintance.  I  was 
myself  at  the  time  a  student  at  the  University  of  Berlin. 

My  first  meeting  with  Professor  Dudley  was  at  Strasburg,  where  he 
had  gone  to  study  under  the  famous  botanist,  De  Bary.  Somewhat  later 
Professor  Dudley  went  to  Berlin,  where  I  was  a  student,  and  I  had  an 
opportunity  of  renewing  the  acquaintance  so  pleasantly  begun  at  Strasburg. 
It  is  seldom  that  I  have  had  the  good  fortune  to  meet  a  man  who  has 
made  upon  me  a  deeper  impression.  The  extraordinarily  fine  quality  of 
Professor  Dudley's  personality  it  is  not  necessary  to  describe  to  those  who 
knew  him.  In  every  sense  of  the  word  he  was  a  gentleman  of  the  finest 
type.  We  little  thought  then  that  it  was  not  going  to  be  many  years  before 
we  should  be  colleagues  in  a  new  university  in  far-away  California,  for  to 
us  then  California  seemed  very  far  away  indeed. 

Just  twenty  years  ago  a  little  band  of  pioneers,  to  which  I  had  the 
great  good  fortune  to  belong,  started  our  University  on  its  career.  Every- 
thing looked  most  promising  and  we  were  all  full  of  enthusiasm  and  hope 
for  the  future.  Two  years  later  Mr.  Stanford  died,  and  the  university 
entered  upon  a  period  of  anxiety  and  privation,  which  was  only  tided  over 
by  the  noble  and  self-sacrificing  devotion  of  Mrs.  Stanford. 

Professor  Dudley  was  called  to  Stanford  as  professor  of  systematic 
botany  in  1892,  but  did  not  come  to  California  until  the  fall  of  1893,  just 
at  the  time  when  the  outlook  was  most  discouraging.  He  naturally  had 
expected  to  have  all  the  necessary  equipment  for  establishing  his  depart- 
ment, and  of  course  nobody  could  have  foreseen  the  unfortunate  condition 
of  things  which  prevailed  at  the  time  he  took  up  his  duties  in  the  autumn 


WILLIAM    RUSSEL    DUDLEY CAMPBELL  13 

of  1893.  Although  it  must  have  been  a  great  disappointment  to  him,  he 
nevertheless  vigorously  set  to  work  to  make  the  best  of  the  situation  and 
for  several  years  before  the  outer  quadrangle  was  built  and  the  present 
botanical  quarters  provided,  he  carried  on  the  work  of  his  department  under 
most  discouraging  conditions.  His  laboratories,  if  such  they  may  have  been 
called,  occupied  the  attic  of  one  of  the  shop  buildings  back  of  the  quad- 
rangle, and  were  very  far  from  satisfactory  either  for  laboratory  or  her- 
barium purposes.  However,  he  began  collecting  assiduously  and  before 
long  the  nucleus  of  the  fine  herbarium  which  he  has  left  to  the  university 
was  brought  together. 

The  flora  of  California  is  a  peculiarly  rich  and  interesting  one  and 
offers  exceptional  opportunities  to  the  student  of  the  problems  of  plant 
distribution.  To  Professor  Dudley,  whose  work  had  been  especially  along 
these  lines,  the  opportunities  for  work  in  his  chosen  field  must  have  been 
very  enticing,  and  doubtless  compensated  in  great  measure  for  some  of  the 
drawbacks  in  other  respects  which  he  must  have  felt  keenly  when  he  came 
to  Stanford.  From  the  time  of  his  arrival,  almost  until  his  death,  he  made 
many  trips  to  all  parts  of  the  state,  collecting  zealously  and  accumulating 
an  invaluable  herbarium  which  remains  to  remind  future  students  of  our 
flora  of  his  tireless  interest  in  his  work. 

Professor  Dudley  paid  especial  attention  to  the  flora  of  the  Sierras, 
and  was  a  recognized  authority  upon  it.  California  is  pre-eminent  in  its 
coniferous  forests,  which  are  unrivaled  in  all  the  world,  and  Professor 
Dudley  soon  became  deeply  absorbed  in  a  study  of  the  distribution  of  these 
magnificent  trees.  A  considerable  number  of  these  are  peculiar  to  Cali- 
fornia and  often  of  very  restricted  range,  like  the  familiar  Monterey  cypress. 
Professor  Dudley  studied  with  especial  care  the  habits  and  distribution 
of  a  beautiful  fir  (Abies  venusta)  which  is  only  known  to  grow  in  the 
Santa  Lucia  range.  He  made  a  number  of  trips  to  this  remote  region 
for  the  purpose  of  studying  this  rarest  of  the  Californian  firs.  His 
acquaintance  however  with  all  of  the  coniferous  trees  was  most  intimate, 
and  he  soon  became  a  recognized  authority  on  the  distribution  of  the  Cali- 
fornian conifers. 

Professor  Dudley's  interest  in  the  study  of  the  distribution  of  the 
forest  trees  naturally  led  him  to  a  study  of  the  problems  of  forestry,  which 
for  the  past  twenty  years  or  so  have  been  arousing  so  much  interest  in  the 
United  States,  and  which  so  deeply  concern  the  welfare  of  the  country. 
As  might  be  expected,  his  sympathies  were  entirely  with  those  who  would 
protect  what  is  left  of  our  magnificent  western  forests  from  the  reckless 
exploitation  of  ignorant  or  unscrupulous  men  who  have  so  devastated  the 


14  DUDLEY    MEMORIAL   VOLUME 

forests  of  the  eastern  states,  and  are  now  threatening  the  great  forests  of 
the  Pacific  Coast.  An  intimate  friend  of  Gifford  Pinchot,  who  has  been 
an  effective  champion  of  the  rights  of  all  the  people  in  our  splendid  forests, 
which  have  been  so  wantonly  devastated,  he  always  stood  for  the  most 
enlightened  views  of  forest  conservation.  The  state  has  never  had  a  more 
devoted  advocate  of  sound  and  modern  methods  in  forestry  than  Professor 
Dudley. 

His  teaching  work  in  the  university,  especially  in  his  later  years,  was 
to  a  great  extent  strongly  influenced  by  his  interest  in  forestry  problems, 
and  the  students  who  were  intending  to  devote  themselves  to  forestry  as  a 
profession  found  in  his  teaching  a  sound  preparation  for  their  future  vocation. 

Professor  Dudley's  interest  in  forestry  was  evinced  in  a  very  practical 
way  through  his  participation  in  the  movement  to  reserve  as  a  state  park 
the  fine  body  of  redwood  timber  in  the  Santa  Cruz  Mountains  known  as 
the  Big  Basin.  Largely  through  his  instrumentality  this  magnificent  body 
of  virgin  redwood  forest  was  bought  by  the  State  as  a  permanent  public 
park.  Until  compelled  by  illness  to  give  up  his  position,  he  served  as 
one  of  the  commissioners  of  the  park,  in  which  to  the  last  he  took  the 
keenest  interest. 

For  many  years  also  Professor  Dudley  was  an  active  and  interested 
member  of  the  Sierra  Club,  and  accompanied  the  club'  in  its  outings  in  the 
Sierras  on  several  occasions.  Those  who  were  fortunate  enough  to  be 
members  of  the  party  and  thus  came  to  know  Professor  Dudley  in  his 
most  congenial  surroundings,  will  always  remember  with  the  keenest  pleasure 
their  associations  with  him  on  those  excursions. 

As  a  teacher  Professor  Dudley  was  devoted  to  the  welfare  of  his 
students,  who  will  bear  witness  to  his  constant  interest  in  their  work  and 
the  unfailing  assistance  always  rendered  them.  Many  students  both  at 
Cornell  and  Stanford  came  under  his  influence,  and  the  long  roll  of  those 
who  have  achieved  success  in  their  work  after  leaving  college  bears  witness 
to  the  success  of  his  labors  as  a  teacher.  At  Cornell,  Professor  Atkinson, 
the  present  head  of  the  department  of  botany,  was  one  of  his  students. 
Professor  Trelease,  the  distinguished  director  of  the  Missouri  Botanical 
Gardens,  which  position  he  recently  resigned,  was  also  a  student  at  Cornell ; 
and  Dr.  Coville,  head  of  the  National  Herbarium  at  Washington,  also 
claims  Professor  Dudley  as  his  teacher.  Many  others,  successful  both  as 
teachers  and  investigators,  look  back  with  pleasure  and  gratitude  to  their 
student  days  in  his  laboratory.  On  our  own  faculty  Professor  Abrams  and 
Mr.  McMurphy  were  both  associated  with  him  as  students  and  colleagues, 
and  are  carrying  on  the  work  which  he  so  well  began. 


WILLIAM    RUSSEL    DUDLEY CAMPBELL  15 

Undoubtedly  Professor  Dudley's  most  important  scientific  work  was  the 
collection  of  the  extensive  herbarium  to  which  he  devoted  so  much  time 
and  labor  during  all  the  years  that  he  spent  in  California.  It  is  doubtful 
whether  any  botanist  had  a  more  intimate  knowledge  of  the  flora  of  Cali- 
fornia than  he,  and  the  great  number  of  specimens  collected  by  him  on 
his  many  botanical  trips  are  now  the  property  of  the  university.  And  the 
Dudley  Herbarium  will  remain  as  a  monument  to  his  devoted  labors  as 
a  student  of  California  plants. 

A  characteristic  California  genus,  Dudleya,  has  been  named  in  his 
honor,  and  will  always  recall  to  botanists  the  name  of  one  of  the  most  devoted 
students  of  the  flora  of  our  state. 


WILLIAM  RUSSEL  DUDLEY* 
By  PRESIDENT  DAVID  STARR  JORDAN 

THE  fact  that  the  writer  has  been  intimately  associated  with  Professor 
Dudley    since    the   day   he   entered   the    freshman   class    at    Cornell 
University,    in   September,    1870,    will   perhaps  excuse   the   personal 
element  in  this  little  sketch. 

The  word  '"instructor"  as  a  technical  term,  describing  a  minor  assistant 
to  a  professor,  had  just  then  been  invented,  and  the  present  writer  had 
just  been  appointed  "instructor  in  botany"  under  Professor  Albert  N. 
Prentiss. 

•  One  day  Professor  Henry  T.  Eddy,  now  of  Minnesota,  brought  to  me 
a  tall,  well-built,  handsome  and  refined  young  man,  older  and  more  mature 
than  most  freshmen,  and  with  more  serious  and  definite  purposes.  Young 
Dudley  had  an  intense  delight  in  outdoor  things  and  especially  in  flowers 
and  birds.  He  wanted  to  be  a  botanist,  and  had  turned  from  old  Yale, 
to  which  as  a  descendant  of  Chittendens,  Griswolds  and  Dudleys  he  would 
naturally  have  gone,  to  new  Cornell,  because  Cornell  offered  special  ad- 
vantages in  science.  For  the  rest  of  my  stay  at  Cornell,  Dudley  was  my 
roommate,  living  in  a  cottage  on  the  hill,  built  by  students  and  termed 
"University  Grove."  In  this  cottage  was  established  the  boarding-club, 
known  later  and  appropriately  as  "The  Struggle  for  Existence,"  and  in 
later  and  more  economical  times  as  the  "Strug." 

In  time  he  was  made  botanical  collector,  and  this  congenial  work  he 
kept  up  until  he  became  my  successor  as  instructor  in  botany.  In  college 
Dudley  was  a  member  of  the  Delta  Upsilon  fraternity,  and  took  an  active 
part  in  holding  this  society  to  the  high  ideals  (AiKtua  yiroOrjKr)}  on  which 
it  was  originally  based.  He  was  also  a  charter  member  in  the  honorary 
scientific  society  of  Sigma  Xi  (STrouSuv  Hw<ov«). 

From  1872  to  1876  he  was  instructor  in  botany  at  Cornell,  his  eminent 
knowledge  of  the  eastern  flora  overbalancing  the  fact  that  at  first,  he  had  not 
yet  received  a  degree.  From  1876  to  1892  he  was  assistant  professor  of 
botany  at  Cornell,  with  a  year's  absence  in  1880,  in  which  he  served  as 
acting  professor  of  biology  in  the  University  of  Indiana,  in  the  absence 
of  the  present  writer,  who  then  held  that  chair. 

In  1892,  Professor  Dudley  became  professor  of  systematic  botany  at 
Stanford  University,  which  position  he  held  until,  in  January,  1911,  failing 


*  Science,  N.  S.,  Vol.  XXXIV,  142-145,  August  4,  1911. 


WILLIAM    RUSSEL    DUDLEY JORDAN  17 

health  caused  his  retirement  on  the  Carnegie  Foundation,  as  professor 
emeritus,  his  work  being  then  taken  by  one  of  his  students,  Associate  Pro- 
fessor LeRoy  Abrams. 

Many  of  the  leading  botanists  of  the  country  have  been  students  of 
Professor  Dudley.  H.  E.  Copeland,  Kellerman,  Lazenby,  Branner  were 
among  his  associates  at  Cornell.  Atkinson  became  his  successor  at  Cornell. 
Abrams,  Cook,  Elmer,  Olssen-Seffer,  Cannon,  Wight,  E.  B.  Copeland, 
E.  G.  Dudley,  Greeley,  Herre,  McMurphy  and  many  others  were  under  his 
tutelage  at  Stanford. 

In  Stanford  University,  Dudley  was  one  of  the  most  respected  as  well 
as  best  beloved  members  of  the  faculty.  No  one  could  come  near  to  him 
without  recognizing  the  extreme  refinement  of  his  nature ;  a  keen  intellect, 
an  untiring  joy  in  his  chosen  work,  and  the  Puritan  conscience  at  its  best, 
with  clear  perceptions  of  his  own  duties  to  himself  and  a  generous  recogni- 
tion of  the  rights  and  the  aspirations  of  others. 

Dudley  entered  with  great  joy  into  the  study  of  the  California  flora. 
He  became  especially  interested  in  the  study  of  trees,  the  evolutionary 
relations  of  forms  and  especially  the  problems  of  geographical  distribution. 
The  conifers  of  California  were  his  special  delight,  and  he  made  many 
field  trips  with  his  students  to  all  parts  of  the  state,  notably  to  the  Sierra 
Nevada  and  the  Sierra  Santa  Lucia.  His  extended  collections  were  presented 
to  Stanford  University,  where  with  the  collections  of  Dr.  Abrams  they  form 
the  major  part  of  the  large  "Dudley  Herbarium." 

A  genus  of  stone-crops,  of  many  species,  abounding  on  the  cliffs  of 
California  and  especially  on  those  which  overhang  the  sea,  was  named 
Dudleya  by  Britton  and  Rose.  Dudleya  pulverulenta  is  one  of  the  most 
conspicuous  plants  in  California  wherever  "sea  and  mountain  meet." 

Dudley  was  instrumental  in  inducing  the  State  of  California  to  pur- 
chase a  forest  of  redwoods  (Sequoia  sempervirens],  that  this,  the  second 
of  California's  giant  trees,  might  be  preserved  in  a  state  of  nature.  Two 
thousand  five  hundred  acres  in  the  "Big  Basin"  of  Santa  Cruz  county  were 
thus  bought  and  established  as  the  "Sempervirens  Park."  For  several  years 
Dudley  served  on  the  board  of  control  of  this  park. 

Of  the  Sierra  Club  of  California,  Dudley  was  a  leading  member  and 
for  some  years  a  director. 

As  an  investigator,  Professor  Dudley  was  persistent  and  accurate,  doing 
his  work  for  the  love  of  it.  A  partial  list  of  his  papers  is  given  below.  A 
large  work  on  the  conifers  of  the  west  was  long  projected,  but  still  exists 
only  in  uncompleted  manuscript. 


18  DUDLEY    MEMORIAL   VOLUME 

Dudley  was  master  of  a  quiet  and  refined  but  effective  English  style. 
He  was  one  of  those  scientific  men,  too  few  I  fear,  who  have  real  love 
for  literature,  and  who  understand  what  poetry  is  and  what  it  is  about. 
In  his  early  days  he  wrote  graceful  verse.  Three  of  his  poems  are  in  print, 
"The  Kaaterskills  as  seen  from  the  Taconics,"  "Sunrise  on  the  Kaaterskill" 
and  "A  Legend  of  the  Lehigh  Valley."  The  last  is  the  story  of  the 
Moravian  settlements  of  "Friedenhutten,  Tents  of  Peace,  and  Gnadenhiitten, 
Tents  of  Grace." 

From  the  first  of  these,  I  quote: 

'Twas  reached  at  last,  with  toiling  long  and  weary 

Taconic's  loftiest  hill; 
Then,  visions  of  all  visions,  stood  uncovered 

The  domes  of  Kaaterskill! 

They  rose  above  the  lesser  hills  as  sovereigns 

Above  the  common  herd; 
They  gathered  then  in  conclave  grand  and  solemn ; 

They  breathed  no  spoken  word. 

But  full  as  anthemed  voices  of  the  ocean 

A  soundless  song  was  borne 
Up  from  those  lips  that  changeless  through  the  ages 

Sang  on  Creation's  morn. 

A  mighty  calm  sits  on  these  silent  summits, 

Time  fades,  as  breath  away, 
O'er  all  in  solemn  oceanic  pulsings 

Deep    flows — Eternity. 

From  '-'A  Legend  of  the  Lehigh  Valley"  I  quote  the  last  verses: 

Full  six  score  years  have  passed  away. 

Still    on    the   silent    summer   morn, 
At  noon's  repose,  or  evening's  gray, 
O'er  Lehigh's  vale  this  dirge  is  borne. 
The  reaper  hears,   on  far-off  hills, 
And  the  traveler  by  the  mountain  rills, 
And  the  fisher  in  the  evening's  chills; 


WILLIAM    RUSSEL    DUDLEY JORDAN  19 

They  hear  and  feel  some  echo  wake 

Of  sorrow  slumbering  long.     A  tear 
Is  shed  for  some  sweet  lost  one's  sake, 

A  tear  that  leaves  life's  stream  more  clear. 
They  bless  the  song  and  them  who  sing ; 
They  feel  the  sympathy  upspring 
That's  born  of  human  suffering. 

The  air  is  full  of  sad-toned  bells 

That  never  cease  their  brazen  toll; 
With  circling  suns  their  pulsing  swells, 
And  in  one  tireless  world-wave  roll. 
But  grateful  unto  sorrow's  ear 
From  the  Lehigh,  far  or  near, 
Comes  this  dirge  so  sweet  and  clear — 
Come  these  human  voices  dear. 

Professor  Dudley's  health  was  good  until  about  three  years  ago,  when 
he  set  out  to  study  the  trees  of  Persia.  In  Egypt  he  was  attacked  by 
a  severe  cold  or  bronchitis  which  ended  in  tuberculosis. 

He  never  married. 


PROFESSOR   DUDLEY'S   WORK  FOR  STANFORD  * 
By  PROFESSOR  LEROY  ABRAMS 

PROFESSOR  William  Russel   Dudley,  who  became  professor  emeritus  of 
botany  at  the  opening  of  the  present  semester,  although  born  in  an  old 
New  England  town  that  has  been  the  home  of  the  Dudley  family  since 
early  colonial  times,  is  essentially  a  pioneer.      Entering  Cornell  University 
with  its  second  freshman  class,  he  remained  in  that  young  institution  after 
graduation,   first  as  instructor  and  later  as  assistant  professor   of   botany, 
until  the  foundation  of  our  own  university,  when  at  the  urgent  request  of 
President  Jordan,  his  college  mate  and  intimate  friend,  he  came  to  Stan- 
ford as  one  of  the  pioneer  professors  at  the  opening  of  its  second  year. 

Of  Professor  Dudley's  experience  at  the  very  beginning  of  work  in 
his  new  field,  and  of  the  arduous  times  during  the  dark  days  that  en- 
gulfed the  university  soon  afterward,  I  have  no  personal  knowledge,  for 
it  was  some  four  or  five  years  after  his  arrival  that  I  came  to  know  him. 
Upon  entering  the  university  I  sought  out  the  department  of  systematic 
botany  with  the  intention  of  carrying  on  some  studies  in  flowering  plants. 
At  that  time  the  twelve  small  buildings  which  form  the  inner  quadrangle, 
and  three  small  shop  buildings  in  the  rear  of  them,  were  the  only  build- 
ings available  for  university  work.  In  my  search  for  the  department  I 
was  directed  to  the  farthest  of  the  shop  buildings,  the  one  situated  just 
back  of  the  new  geology  building,  where  I  was  told  that  I  would  find 
Professor  Dudley  on  the  second  floor.  And  here  I  did  find  him,  tucked 
away  in  one  end  of  a  loft,  in  a  single  room,  one  corner  of  which  had 
been  partitioned  off  as  an  office.  In  a  quiet,  reserved  manner  he  talked 
over  my  work;  then  he  took  me  into  the  main  room  to  select  a  table  and 
material  for  study.  It  was  a  curious-  room,  this  "laboratory,"  perched 
high  amid  the  rafters.  Three  huge  beams  ran  lengthwise  of  it  a  good 
hurdling  distance  apart,  but  about  five  feet  and  a  half  from  the  floor. 
With  an  apologetic  smile,  he  warned  me  of  these  as  he  calmly  ducked  under 
the  first.  The  table  was  soon  selected  and  my  initial  study  outlined.  Day 
by  day,  throughout  the  course,  as  he  went  from  student  to  student  directing 
their  studies,  he  patiently  dodged  those  formidable  beams. 

For  ten  years  this  man,  one  of  America's  foremost  teachers  of  botany, 
conducted  his  classes  under  such  handicaps.  Yet  with  these  great  obstacles 
constantly  checking  the  normal  growth  and  development  of  his  cherished 
plans,  he  labored  on  incessantly;  his  quiet,  dignified,  courteous  manner, 


*From  the  Stanford  Alumnus,  Vol.  XII,  No.  6,  pp.  165-166,  February,  1911. 


PROFESSOR  DUDLEY'S  WORK  FOR  STANFORD  —  ABRAMS  21 

his  thoroughness  and  enthusiasm  in  his  work,  his  broad  interests  and  scholarly 
attainments  moulding  the  lives  of  his  students.  For  none  can  come  under 
his  influence  without,  at  least  unconsciously,  acquiring  higher  ideals  and 
more  serious  purposes. 

During  the  summer  vacations  the  pursuit  of  his  botanical  studies  took 
him  into  the  mountains  and  forested  areas  of  the  state,  where  he  was  con- 
stantly confronted  with  the  great  and  shameless  waste  of  our  forest  resources. 
He  thus  became  one  of  the  pioneers  in  the  movement  toward  conservation, 
and  rendered  valuable  service  to  the  state  and  nation  through  suggestions 
and  advice  to  the  Forest  Service  and  other  authorities.  The  establishment 
of  the  California  Redwood  Park,  a  beautiful  tract  of  forested  land  in  our 
neighboring  mountains,  set  aside  by  the  state  primarily  for  the  purpose 
of  preserving  a  forest  of  the  coast  redwood  in  its  primitive  conditions,  was 
accomplished  largely  through  his  efforts.  And  as  secretary  of  the  first  park 
commission  he  labored  for  its  betterment  and  the  establishment  of  a  per- 
manent policy  in  its  management. 

But  Professor  Dudley  saw  that  if  the  conserving  of  our  forests  was 
to  be  placed  on  an  intelligent  and  permanent  basis  it  was  essential  that 
young  men  be  trained  for  the  work,  and  that  the  people  of  the  states 
where  the  forests  abound  be  educated  to  the  necessity  of  scientific  forestry; 
he  saw  that  fully  nine-tenths  of  the  nation's  forests  lay  west  of  the  con- 
tinental divide,  yet  in  all  this  region  not  one  of  the  educational  institutions 
was  training  men  for  the  scientific  management  of  this  vast  wealth.  He 
therefore  directed  his  energies  toward  the  establishment  of  courses  in  forestry 
at  Stanford.  For  a  number  of  years  he  planned  toward  this  end,  and  finally, 
just  as  success  seemed  probable,  the  fateful  April  18th  wiped  out  every 
promising  hope  of  immediate  realization.  Soon  afterward  he  contracted  a 
serious  illness  which  left  him  physically  weakened.  This  hampered  his 
work,  but  not  his  enthusiasm,  and  he  is  now  retiring  from  the  regular 
routine  departmental  duties  in  the  hope  that  he  may  regain  his  health 
sufficiently  to  complete  his  research  studies  on  the  western  flora. 

Professor  Dudley's  students  and  his  many  other  friends  who  have 
known  and  followed  his  courageous  and  uncomplaining  struggle  against 
disheartening  obstacles  hope  that  he  may  not  only  live  to  complete  his 
own  studies,  but  that  he  may  yet  see  young  men  trained  at  Stanford  for 
the  scientific  management  of  the  vast  forests  of  the  West. 


PROFESSOR  WILLIAM  RUSSEL  DUDLEY* 
By  PROFESSOR  GEORGE  JAMES  PEIRCE 

WILLIAM  RUSSEL  DUDLEY,  professor  of  systematic  botany  in  Leland 
Stanford  Junior  University  from  1892  to  1911,  died  on  June  4,  1911, 
at  the  age  of  sixty-two.  By  ancestry  and  place  of  birth  a  New 
Englander,  a  graduate  and  for  twenty  years  -a  member  of  the  botanical  staff 
of  Cornell  University,  a  student  of  De  Bary's  for  a  time  in  Strasburg,  he 
brought  to  California  the  mature  powers  of  an  enthusiastic  student  and 
sympathetic  lover  of  nature,  the  ripe  scholarship  and  the  winning  personality 
of  the  inspiring  teacher.  At  home  in  the  laboratory,  he  was  still  more 
strikingly  the  gracious  host  when  he  was  with  students  and  other  friends 
out  of  doors,  in  the  fields  and  woods  and  mountain  forests. 

He  knew  the  forests  of  middle  California  as  no  one  else;  his  acquaint- 
ance was  with  individual  trees,  as  his  collection  of  tree  portraits  testifies. 
And  his  studies  of  their  geographical  distribution,  following  and  amplify- 
ing the  earlier  studies  of  Asa  Gray  and  others,  gave  his  knowledge  a  degree 
of  accuracy  and  detail,  as  well  as  breadth,  which  was  very  precious.  It 
is  to  be  hoped  that  his  notes  and  other  manuscripts  are  in  such  condition 
that  his  associates  and  successor  can  give  them  to  the  world. 

Professor  Dudley's  nature  was  so  sensitive,  his  perceptions  so  fine,  and 
his  ideals  so  high,  that  he  could  but  rarely  bring  himself  to  publish  what 
he  knew.  He  wished  always  to  add  to  and  improve  what  he  had  learned 
and  written.  Thus  the  botanical  world  had  little  opportunity  to  know  his 
accomplishments  and  achievements. 

Besides  the  young  men  and  women  whose  lives  he  has  enriched,  and 
the  Forest  Service  which  he  long  assisted  in  various  ways,  he  contributed 
to  the  great  gift  to  California  and  the  nation  which  the  state  and  national 
forests  of  California  constitute.  The  "Big  Basin  Park,"  the  property  of 
the  state,  will  preserve  to  all  time  a  part  of  the  natural  redwood  forest 
of  the  Santa  Cruz  mountains.  Professor  Dudley  assisted  in  securing  and 
preserving  as  a  state  park  this  part  of  the  virgin  forest  of  Sequoia  semper- 
virens.  It  was  his  interest  too  which  stimulated  and  directed  the  federal 
authorities  in  the  selection  of  others  of  the  mountain  forests  of  California 
as  national  forests. 

Of  courtly  manner,  cultivated  as  well  as  educated,  of  ripe  scholarship 
and  rich  in  the  knowledge  of  nature,  he  was  a  lovable  and  elevating  associate, 
an  inspiring  teacher,  a  devoted  man  of  science,  an  honor  to  Stanford  Uni- 
versity of  which  he  was  an  honored  member. 


From  The  Plant  World,  Vol.  XIV,  No.  8,  pp.  200-202,  August,  1911. 


PROFESSOR  WILLIAM   RUSSEL  DUDLEY* 
By  JARED  TREMAN  NEWMAN 

ONE  of  the  purest  and  noblest  souls — such  as  one  is  fortunate  to  come 
close  to  even  once  or  twice  in  a  lifetime — passed  to  the  life  beyond 
yesterday  afternoon. 

Professor  Dudley  was  a  prominent  scientist,  "one  of  America's  fore- 
most teachers  of  botany,  one  of  the  pioneers  in  the  movement  toward 
conservation,"  largely  instrumental  in  the  establishment  of  the  California 
Redwood  Park,  and  the  secretary  of  the  park  commission;  yet,  it  is  not  of 
these,  nor  of  his  other  scientific  attainments  or  accomplishments,  that  we 
think  chiefly  at  this  time. 

Of  fine  New  England  stock,  cultured,  with  a  refinement  that  was 
genuine  all  the  way  thorough,  doing  splendid  work  in  his  chosen  profession 
and  capable  of  making  a  great  name  for  himself,  his  best  service  to  the  world 
was  in  imparting  to  other  men  higher  aspirations  and  nobler  ideals. 

Far  back  in  the  early  days  at  Cornell,  there  was  a  little  coterie  of  men 
gathered  in  close  association.  It  included  Jordan  and  Branner  and  Nichols 
and  Gage  and  Fairchild  and  Kellerman  and  many  others  who  have  deservedly 
come  to  high  position.  Among  them  all  there  was  none  of  finer  instincts 
or  more  lovable  character  than  Dudley. 

For  many  years  after  his  graduation  at  Cornell,  and  while  he  remained 
a  teacher  there,  he  was  the  guiding  and  inspiring  genius  of  successive  groups 
of  young  men.  Some  were  taking  his  work.  Others  were  attracted  to  him 
by  his  rare  personality.  Still  others  he  sought  out.  What  he  imparted  to 
them,  and  to  all  who  came  close  to  him,  was  something  of  priceless  value. 
It  was  the  very  soul  of  the  man.  He  withheld  nothing.  Absolutely  un- 
calculating  in  his  unselfishness,  so  pure  that  impurity  could  not  be  thought 
of  in  his  presence,  a  lover  of  nature  and  nature's  God,  his  influence 
was  constantly  ennobling.  Like  many  noble  souls,  he  was  peculiarly  sensi- 
tive. He  was  hurt  often  when  no  hurt  was  intended.  He  was  often 
melancholy,  sometimes  almost  morbid.  It  has  always  seemed  so  strange 
that  one  who  gave  so  much  and  so  constantly  should  not  be  always  happy. 
Perhaps  he  made  up  for  it  in  the  intensity  of  his  joys.  While  he  was  often 
misundertood  and  while  the  number  of  persons  who  came  close  to  him  was 
not  relatively  large,  yet  few  men  have  merited,  or  have  known,  in  so  large 
a  degree,  the  love  of  their  fellows. 


From  The  Palo  Alto  Times,  June  5,  1911. 


24  DUDLEY    MEMORIAL    VOLUME 

A  lover  of  truth  and  imbued  with  the  scientific  spirit,  he  might  have 
become  more  famous  had  he  spent  more  time  in  research  and  in  publishing 
the  results;  but  his  principal  work  is  of  the  kind  that  lives  in  the  hearts 
of  living  men,  and  goes  on,  and  will  continue  to  go  on,  in  a  generation  of 
workers  who  owe  to  him  the  touch  that  makes  their  work  worth  while. 


WILLIAM   RUSSEL   DUDLEY 

[Read  before  the  Stanford  Alumni  Association  at  Washington,  D.  C, 
November    11,    1911.] 

By  WILLIAM  FRANKLIN  WIGHT 

DURING  the  early  summer  one  of  Stanford's  most  lovable  teachers 
closed  his  life's  work  and  found  that  last  long  rest  which  must  come 
to  us  all.  I  wish  therefore  to-night  to  pay  a  brief  tribute  to  the 
memory  of  Professor  William  Russel  Dudley.  His  kindly  feeling  and  in- 
terest in  his  students  made  him  loved  by  them  all,  and  he  possessed  that 
indescribable  quality  in  a  teacher  that  without  thought  and  without  effort 
instantly  arouses  enthusiasm  in  the  laboratory  and  in  the  classroom.  He 
was  an  unusual  teacher,  and  it  is  a  sad  thought  to  realize  that  years  before 
the  allotted  time  of  life  his  voice  will  be  heard  no  more  in  the  classroom 
and  his  charming  manner  will  be  unknown  to  the  students  who  shall  fill 
the  halls  of  Stanford. 

It  was  my  fortune  to  be  with  him  on  the  last  day.  I  had  visited  him 
a  few  weeks  earlier,  and  then  he  was  hopeful  that  there  might  still  be  left 
to  him  a  few  years  in  which  to  complete  the  botanical  work  that  he  had 
begun  almost  immediately  on  coming  to  California.  Nevertheless,  those  who 
saw  him  knew  that  it  was  even  then  too  late — that  the  end  must  soon  come. 
It  was  therefore  with  a  sad  heart  that  I  went  on  the  morning  of  June  4th 
to  pay  a  last  visit  to  my  friend  and  teacher. 

From  the  balcony  where  he  lay  in  the  cottage  at  Los  Altos  one  could 
look  across  the  valley  to  the  Mount  Hamilton  range  bathed  in  sunlight,  and 
view  the  glory  of  a  California  landscape.  The  air  was  crisp  and  full  of 
life  to  the  strong.  It  was  indeed  a  beautiful  day  in  which  to  live,  but 
there  with  the  vision  of  nature  he  loved  so  well  before  him,  now  too  far 
away  for  his  eyes  to  see,  in  the  midst  of  a  few  friends,  he  calmly  awaited 
the  end. 

It  is  however  of  other  days  that  we  would  keep  the  memories  fresh. 
We  would  rather  remember  him  strong  and  enjoying  the  activities  of  a  busy 
life.  And  I  think  he  took  keen  pleasure  in  all  his  work,  for  he  appeared 
.to  go  through  each  year  at  the  university  with  an  enthusiasm  equal  to  that 
we  should  expect  if  the  studies  and  discoveries  of  the  laboratory  were  as 
new  to  him  as  to  the  student.  But  it  was  on  long  tramps  in  the  mountains, 
in  the  solitude  and  grandeur  of  the  redwood  forest,  that  one  really  began 
to  appreciate  the  fineness  of  the  man,  to  know  how  much  he  saw  in  mountain 
and  forest,  and  how  much  he  loved  nature  in  sunshine  and  in  storm.  At 


26  DUDLEY    MEMORIAL    VOLUME 

fifty  years  there  was  the  freshness  and  joy  of  youth  in  botanical  exploration. 
It  was  when  on  such  walks  too  that  one  came  to  know  the  fullness  of  his 
knowledge  and  how  perfect  was  his  memory,  as  every  species  was  recognized 
and  its  distribution  or  other  fact  of  interest  was  related. 

In  many  ways  his  life  at  Stanford  was  a  disappointment.  He  felt  the 
burden  of  the  years  of  financial  stress  through  which  the  university  passed 
more  than  was  his  share,  and  very  often  supplied  from  his  own  purse  the 
necessary  materials  for  the  laboratory.  The  hopes  and  ideals  he  had  for 
the  development  of  botanical  science  he  could  not  live  to  see  realized.  But 
whatever  was  lacking  in  appropriate  rooms  and  equipment  was  more  than 
compensated  for  in  the  ability  and  spirit  of  the  teacher.  He  lived  in  his 
work  and  for  his  students.  His  time  and  energy  were  so  very  largely  oc- 
cupied in  their  interest  that  he  published  little,  and  this  is  the  regret  of  all 
who  realize  the  high  scientific  ideal  which  guided  him  in  his  work,  and  who 
appreciate  the  charm  of  his  literary  style.  His  Flora  of  Ithaca  and  of  the 
Wyoming  Valley  will  be  regarded  as  classics  and  as  models  of  their  kind 
for  many  years  to  come.  For  some  it  is  not  given  to  publish  much — it  is 
theirs  to  write  in  the  hearts  and  minds  of  men  and  women,  an  influence  as 
enduring  perhaps  as  that  of  printed  books. 

I  never  heard  him  speak  ill  of  any  person  but  once,  and  then  he  did 
it  deliberately,  reluctantly,  and  as  though  he  felt  it  a  painful  duty.  It  was 
his  habit  to  see  the  good  qualities  in  mankind  and  he  did  it  naturally  and 
without  effort. 

I  trust  that  so  long  as  modesty,  thoughtfulness,  and  a  kindly  spirit 
are  regarded  as  evidences  of  a  fine  character,  that  so  long  the  memory  of 
Professor  William  Russel  Dudley  will  live  at  Stanford  University. 


PUBLICATIONS  OF  WILLIAM   RUSSEL  DUDLEY 

First  Steps  in  the  Study  of  Botany.     Educationist,  2:  7-10.     1880. 

Leafy  Berries  in  Mitchella  repens.     Bull.  Torr.  Bot.  Club,  10 :  1-3.     1883. 

An  Abnormal  Orchid.  Abstract  of  a  paper  read  before  the  American 
Association  for  the  Advancement  of  Science.  Proc.  Amer.  Assoc.  Adv. 
Sci.  32:  30.  1883. 

Sketch  of  Curtis.     Jour.  Mycology,  2:  54-59.     1886. 

Elias   Magnus   Fries.     Jour.   Mycology,   2:   91-94.     1886. 

Charles  Christopher   Frost.     Jour.   Mycology,  2:    114-118.     1886. 

The  Cayuga  Flora,  Part  1.  A  catalogue  of  the  Phaenogamia  growing 
without  cultivation  in  the  Cayuga  Lake  Basin.  Bull.  Cornell  Univ. 
Sci.  2:  XXX,  1-132.  1886. 

A  Preliminary  List  of  the  Vascular  Plants  of  the  Lackawanna  and  Wyoming 
Valleys.  Proc.  and  Coll.  Lackawanna  Inst.  Hist,  and  Sci.  1:  32-112. 
1887. 

Strasburg  and  its  Botanical   Laboratory.     Bot.   Gaz.     13:  305-311.     1888. 

The  Death  of   DeBary.     Bot.   Gaz.     13:  64-65.     1888. 

Report  of  the  Cryptogamic  Botanist.  Rep.  Cornell  Agri.  Exp.  Sta.  1-3. 
1888-1890. 

The  Strawberry  Leaf-Blight.  Rep.  Cornell  Agri.  Exp.  Sta.  2:  171-182. 
1889. 

Another  Disease  of  the  Strawberry.  Rep.  Cornell  Agri.  Exp.  Sta.  2 : 
182-183.  1889. 

Anthracnose  of  Currants.     Rep.  Cornell  Agri.  Exp.  Sta.    2:  196-198.    1889. 

Leaf-Blight  of  Quince  and  Pear.  Rep.  Cornell  Agri.  Exp.  Sta.  2:  198-199. 
1889. 

The  Onion  Mold.     Rep.  Cornell  Agri.  Exp.  Sta.     2:  193-196.     1889. 

The  Clover  Rust.  J.  K.  Howell.  Footnote  by  W.  R.  Dudley.  Rep.  Cor- 
nell Agri.  Exp.  Sta.  3:  129-139.  1890. 

The  Hollyhock  Rust.     Rep.  Cornell  Agri.   Exp.  Sta.     3:   154-155.     1890. 

Flora  of  the  Lackawanna  and  Wyoming  Valleys;  a  catalogue  of  the  flower- 
ing plants  and  vascular  Cryptogams  found  in  and  near  Lackawanna 
and  Wyoming  Valleys.  With  Charles  O.  Thurston.  Wilkesbarre,  Pa. 
xv,  96.  1892. 

The  Genus  Phyllospadix.  Wilder  Quarter- Century  Book,  Comstock  Pub- 
lishing Co.,  Ithaca,  N.  Y.  403-420.  1893. 

Botanical  Notes.     Bull.   Torr.   Bot.  Club,  20:   169-170.     1893. 

Phyllospadix;  its  Systematic  Characters  and  Distribution.  Zoe,  4:  381-385. 
1894. 


28  DUDLEY    MEMORIAL   VOLUME 

A  Laboratory  Manual  of  Plant  Histology.  With  Mason  B.  Thomas,  Craw- 
fordsville,  Ind.  viii,  115.  1894. 

Forest  Reservations;  With  a  Report  on  the  Sierra  Reservation,  California. 
Sierra  Club  Bull.  1:  254-267.  1896. 

The  Kaweah  Group.     Sierra  Club  Bull.     5:185-191.     1898. 

Forestry  Notes,  editor  of.     Sierra  Club  Bull.     2-7.     1898-1910. 

A  Short  Account  of  the  Big  Trees  of  California,  (one  of  the  collaborators). 
Bull.  U.  S.  Dept.  Agri.  Div.  Forestry,  No.  28;  1-30.  1900. 

The  Big  Trees  of  California.     Forester,  6:  206-210.     1900. 

Lumbering  in  the  Sequoia  National  Park.     Forester,  6:  293-295.     1900. 

Zonal  Distribution  of  Trees  and  Shrubs  in  the  Southern  Sierra.  Sierra 
Club  Bull.  3:  298-312.  1901. 

Big  Basin  Redwood  Park.     Forester,  7:  157-164.     1901. 

A  Notable  California  Fir.     Forestry  and  Irrigation,  8:   193-198.     1902. 

Trees  along  the  Tulare  Trails.     Sierra  Club  Bull.     4:   153-156.     1902. 

Trees  of  Southern  California.  Los  Angeles  Saturday  Post,  5-6.  June  7- 
August  2,  1902. 

Near  the  Kern's  Grand  Canon.     Sierra  Club  Bull.     4:  301-307.     1903. 

Notes  on  California's  Uredineas  and  Descriptions  of  New  Species.  (With 
C.  H.  Thompson.)  Jour.  Mycology,  10:  52-55.  1904. 

The  Vitality  of  Sequoia  Gigantea.  Read  by  invitation  before  the  California 
Alumni  Association  of  Columbia  University.  Privately  printed  by  mem- 
bers of  the  association.  San  Francisco,  16.  1905.  • 

Concerning  the  Vitality  of  Sequoia  Sempervirens.  Palo  Alto  Times,  March. 
17,  1908. 


CORNELL   UNIVERSITY   PUPILS   OF 
PROFESSOR  DUDLEY 

ARTHUR,  JOSEPH  CHARLES;  B.S.,  M.S.,  Sc.D. 

Professor    of   Vegetable    Physiology    and    Pathology,    Purdue    University, 

Lafayette. 
ATKINSON,  GEORGE  FRANCIS;  Ph.B. 

Professor  of  Botany,  Cornell  University,  Ithaca. 
BRAY,  WILLIAM  L.;  A.B.  A.M.,  Ph.D. 

Professor  of  Botany,  Syracuse  University,  Syracuse. 
CHESTER,  FREDERICK  DIXON;  B.S.,  M.S. 

Bacteriologist,    (formerly)    Director,    State   Bacteriological   Laboratory   of 

Delaware. 
CORBETT,  LEE  CLEVELAND;  B.S.,  M.S. 

Horticulturist,  Department  of  Agriculture,  Washington. 
COVILLE,  FREDERICK  VERNON;  A.B. 

Botanist,  Department  of  Agriculture,  Washington. 
CRAIG,  MOSES;  M.S. 

Missouri  Botanical  Garden,  St.  Louis. 
DENSMORE,  HIRAM  DELOS;  A.B.,  A.M. 

Professor  of  Botany,  Beloit  College. 
DURAND,  ELIAS  JUDAH;  A.B.,  Sc.D. 

Assistant  Professor  of  Botany,  University  of  Missouri,  Columbia. 
GREGORY,  EMILY  LORINA. 

Late  Professor  of  Botany,  Barnard  College,  Columbia  University,  New  York. 
HENDERSON,  Louis  FOURNIQUET;  Ph.B. 

Formerly  Professor  of  Botany,  University  of  Idaho,  Moscow. 
HICKS,  HENRY;  B.S. 

Nurseryman,  Westbury,  Long  Island. 
HOFFMAN,  HARRY  NATT;  B.Agr. 

Nurseryman,  Elmira,  New  York. 
HOUGH,  ROMYN  BECK;  A.B. 

Dendrologist,  Lowville,  New  York. 
HOWELL,  JENNY  KIRK;  Ph.B.,  M.S. 

Teacher,  Plainfield,  New  Jersey. 
KELLERMAN,  WILLIAM  ASHBROOK;  B.S.,  Ph.D. 

Late  Professor  of  Botany,  Ohio  State  University,  Columbus. 
LAZENBY,  WILLIAM  RANE;  B.Agr.,  M.Agr. 

Professor  of  Forestry,  Ohio  State  University,  Columbus. 
MOORE,  VERANUS  ALVA;  B.S.,  M.D. 

Director,  New  York  State  Veterinary  College,  Cornell  University,  Ithaca. 
NORRIS,  HARRY  WALDO;  A.B.,  A.M. 

Professor  of  Zoology,  Grinnell  College,  Grinnell,  Iowa. 
PORTER,  EDNA. 

Botanical  Gardens,  Buffalo,  New  York. 

ROWLEE,    WlLLARD    WlNFIELD;    B.L.,  Sc.D. 

Professor  of  Botany,  Cornell  University,  Ithaca. 
SCHRENK,  HERMANN  VON;  B.S.,  A.M.,  Ph.D. 

Consulting  Timber  Engineer,  Plant  Pathologist,  St.  Louis. 
SMITH,  THEOBALD;  Ph.D.,  M.  D.,  A.M.,  LL.D. 

Professor  of  Comparative  Pathology,  Harvard  Medical  School,  Boston. 
SNOW,  JULIA  WARNER;  B.S.,  M.S.,  Ph.D. 

Associate  Professor  of  Botany,  Smith  College,  Northampton. 
THOMAS,  MASON  BLANCHARD;  B.S. 

Late  Professor  of  Botany,  Wabash  College,  Crawfordville. 
TRELEASE,  WILLIAM;  B.S.,  Sc.D.,  LL.D. 

Formerly  Director,  Missouri  Botanical  Garden,  St.  Louis. 
WHITE,  CHARLES  DAVID;  B.S. 

Geologist,  Geological  Survey,  Washington. 
YATABE,  RYOKICHI;  B.S. 

Late  Professor  of  Botany,  Imperial  University,  Tokyo,  Japan. 


STANFORD    UNIVERSITY    PUPILS    OF 
PROFESSOR  DUDLEY 

ABRAMS,  LE.ROY;  M.A.,  Ph.D. 

Associate  Professor  of  Botany,  Stanford  University,  California. 
ALDRICH,  JOHN  MERTON;  Ph.D. 

Professor  of  Biology,  University  of  Idaho,  Moscow,  Idaho. 
ANDERSON,  MALCOLM  PLAYFAIR;  B.A. 

Naturalist,  British  Museum,  London,  England. 
ATKINSON,  WILLIAM  SACKSTON;  B.A. 

Scientific  Illustrator,  Stanford  University,  California. 
BAKER,  CHARLES  FULLER;  M.A. 

Professor  of  Zoology,  Pomona  College,  Claremont,  California. 
BELL,  RUBY  GREEN  (MRS.);  M.A. 

M.  Albert  W.  Smith,  Ithaca,  New  York. 
BERRY,  SAMUEL  STILLMAN;  M.A.,  Ph.D. 

Graduate  Student,  Stanford  University,  California. 
BILLINGS,  FREDERICK  HORATIO;  M.A.,  Ph.D. 

Associate  Professor  of  Botany  and   Bacteriology,   University  of  Kansas, 

Lawrence,  Kansas. 
BORING,  ORAMANDA. 

Teacher  of  Botany  and  Physiology,  Stockton  High  School,  Stockton,  Cali- 
fornia. 
BRYAN,  MARY  KATHERINE;  B.A. 

Scientific  Assistant,  United  States  Department  of  Agriculture,  Washington, 

D.  C. 
BURKE,  CHARLES  VICTOR;  M.A.,  Ph.D. 

Graduate  Student,  Stanford  University,  California. 
BURNHAM,  STEWART  HENRY. 

Botanist,  New  York  State  Museum,  Albany,  New  York. 
CANNON,  WILLIAM  AUSTIN;  M.A.,  Ph.D. 

Staff  Member,  Desert  Laboratory,  Carnegie  Institution,  Tucson,  Arizona. 
CHAPMAN,  BERTHA  LOUISE;  M.A. 

Lecturer,  Writer,  Teacher  of  Private  Nature  Study  Classes,  Kansas  City, 

Missouri,  M.  Vernon  Mosher  Cady. 
CHASE,  RAYMOND  EUGENE. 

Principal,  Reno  High  School,  Reno,  Nevada. 
COOK,  MELVILLE  THURSTON;  M.A.,  Ph.D. 

Plant    Pathologist,    Delaware   Agricultural    Experiment    Station,  Newark, 

Delaware. 
COOPER,  ALICE  CECILIA;  M.A. 

Teacher,  Los  Angeles  Polytechnic  High  School,  Los  Angeles,  California. 
COPELAND,  EDWIN  BINGHAM;  Ph.D. 

Dean  and  Superintendent,  College  of  Agriculture,  University  of  Philippines, 

Los  Bafios,  P.  I. 
COUCH,  MARY  JUANITA;  B.A. 

Teacher,  Vacaville,  California. 
CRAVENS,  MARY  RUHAMA;  M.A. 

Teacher,  Sacramento,  California. 
DOANE,  RENNE  WILBUR;    B.A. 

Assistant  Professor  of  Entomology,  Stanford  University,  California. 
DUDLEY,  ERNEST  GRISWOLD;  B.A. 

Assistant  Supervisor,  Sierra  National  Forest,  North  Fork,  California. 
ELMER,  ADOLPH  DANIEL  EDWARD;  M.A. 

Botanist  and  Botanical  Collector,  Publisher  of  Botanical  Leaflets,  Manila, 

P.  I. 
FISHER,  WALTER  KENDRICK;  M.A.,  Ph.D. 

Assistant  Professor  of  Zoology,  Stanford  University,  California. 
FULLAWAY,  DAVID  TIMMINS;  M.A. 

Entomologist,  United  States  Agricultural  Experiment  Station,  Honolulu, 
T.  H. 


STANFORD    UNIVERSITY    PUPILS    OF    PROFESSOR    DUDLEY  31 

GEIS,  HELEN  DUDU;  B.A. 

Teacher,  Los  Angeles  Polytechnic  High  School,  Los  Angeles,  California. 
GREELEY,  ARTHUR  WHITE;  M.A. 

Late  Professor  of  Zoology,  Washington  University,  St.  Louis. 
GRINNELL,  JOSEPH;  M.A. 

Director,  Museum  of  Vertebrate  Zoology,  University  of  California,  Berkeley, 

California. 
HALSEY,  STELLA  DUFFIELD;  B.A. 

Teacher,  San  Diego  High  School,  San  Diego,  California. 
HELLER,  EDMUND;  B.A. 

Naturalist  of  the  Roosevelt  Expedition  to  Africa,  1909-1910,  United  States 

National  Museum,  Washington,  D.  C. 
HERRE,  ALBERT  WILLIAM  CHRISTIAN  THEODORE;  M.A.,  Ph.D. 

Vice- Principal,  Fruitvale  High  School,  Fruit  vale,  California. 
HIGLEY,  ROSE  MIRIAM;  M.A. 

Teacher,  San  Rafael  High  School,  San  Rafael,  California. 
HOLMAN,  RICHARD  MORRIS;  B.A. 

Instructor  in  Botany,  College  of  Agriculture,  University  of  the  Philippines, 

Los  Bafios,  P.  I. 
HUMPHREY,  HARRY  BAKER;  Ph.D. 

Assistant  Professor  of  Botany,  Washington  State  College,  Pullman,  Wash- 
ington. 
HUMPHREY,  OLIVE  AGATHA  MEALEY  (MRS.);  B.S. 

Pullman,  Washington. 
JENKINS,  HUBERT  OLIVER;  B.A. 

Health  Officer,  Palo  Alto,  California. 
KIMURA,  TOKUZA;  B.A. 

Teacher  of  Biology,  Sotokufu  Chugakko,  Formosa,  Japan. 
KNOCHE,  EDWARD  Louis  HERMAN;  B.A. 

Student  of  Botany,  traveling  in  Europe  until  1912. 
KROECK,   Louis  SAMUEL;  M.A. 

Teacher,  College  of  the  Pacific,  San  Jose,  California. 
KUWANA,  SHINKAI  INOKICHI;  M.A. 

Imperial  Agricultural  Experiment  Station,  Tokyo,  Japan. 
McCRACKEN,  MARY  ISABEL;  M.A.,  Ph.D. 

Assistant  Professor  of  Entomology  and  Bionomics,  Stanford  University 

California. 
MCGREGOR,  ERNEST  ALEXANDER;  M.A. 

United  States  Bureau  of  Entomology,  Washington,  D.  C. 
MACKAY,  MINNIE  LAURIE;  B.A. 

Teacher,  Santa  Clara  High  School,  Santa  Clara,  California. 
MCMURPHY,  JAMES  IRA  WILSON;  M.A. 

Instructor  in  Botany,  Stanford  University,  California. 
MILLER,  JOHN  MARTIN. 

Forest  Assistant,  United  States  Forest  Service. 
MORRIS,  CHARLES  SHOEMAKER;  M.A. 

Teacher  of  Botany  and  Zoology,  Palo  Alto  High  School,  Palo  Alto,  Cali- 
fornia. 
MORRIS,  EARL  LEONARD. 

County  Entomologist,  Santa  Clara  County,  and  Field  Assistant,  University 

of  California,  San  Jose,  California. 
NOHARA,  SHIGEROKU. 

Assistant   in    Botany,    Agricultural    College,  Imperial    University,  Tokyo, 

Japan. 
OLSSON-SEFFER,  PEHR  HJALMAR;  Ph.D. 

Deceased. 
PEMBERTON,  CYRIL  E. ;  B.A. 

Field  Agent,  United  States  Bureau  of  Entomology,  Lindsay,  California. 
PEMBERTON,  JOHN  ROTHWELL;  B.A. 

Geologist,  Argentine  Republic,  South  America. 
PETERSON,  ELSA;  B.A. 

Teacher,  Normal  School,  Honolulu,  T.  H. 


32  DUDLEY    MEMORIAL   VOLUME 

PRICE,  WILLIAM  WIGHTMAN;  M.A. 

Fallen  Leaf,  Lake  Tahoe,  California. 
RANDALL,  JOSEPHINE  Dows;  B.A. 

Sometime  Assistant  in  Botany,  Stanford  University,  California. 
RANDOLPH,  FLORA  ALBERTINE;  M.A. 

Principal  Miss  Randolph's  School,  Berkeley,  California. 
ROSE,  JESSIE  PERKINS;  M.A. 

Teacher,  Fort  Klamath,  Oregon. 
RUST,  EVERETT  WINDER;  B.A. 

Assistant  Government  Entomologist,  Lima,  Peru. 
SCOFIELD,  WILLIAM  LAUNCELOT;  B.A. 

Student,  Yale  Forest  School,  Yale  University,  New  Haven,  Connecticut. 
SEALE,  ALVIN;  B.A. 

Chief  of  Department  of  Fisheries,  Bureau  of  Science,  Manila,  P.  I. 
SHAFER,  GEORGE  DANIEL;  M.A.,  Ph.D. 

Engaged  in  research  work  in  Entomology,  East  Lansing,  Michigan. 
SHERFY,  SAMUEL  HASH;  B.A. 

Mount  Morris,  Illinois. 
SHOW,  STUART  BEVIER;  B.A.,  M.F. 

Forest  Assistant,  United  States  Forest  Service. 
SMITH,  CHARLES  PIPER;  M.A. 

Assistant  Professor  of  Botany,  Utah  Agricultural  College,  Logan,  Utah. 
SNODGRASS,  ROBERT  EVANS. 

Artist,  New  York  City. 
STARK,  WILLIAM  HARVEY. 

Nurseryman,  Stark  City,  Missouri. 
STOKES,  SUSAN  GABRIELLA;  M.A. 

Teacher,  Orange  Union  High  School,  Orange,  California. 
SWENSON,  JOHN  CANUTE;  B.A. 

Professor  of  History  and  Economics  and  Dean  of  the  College,   Brigham 

Young  University,  Provo,  Utah. 
THOMPSON,  CHARLES  HENRY. 

In  charge  Department  of  Succulent  Plants,   Missouri   Botanical   Garden, 

St.  Louis,  Missouri. 
TRACY,  HIRAM  HARWOOD. 
WIGHT,  WILLIAM  FRANKLIN;  M.A. 

United  States  Department  of  Agriculture,  Bureau  of  Plant  Industry,  Wash- 
ington, D.  C. 
WILLIAMS,  FLORENCE;  B.A. 

Sometime  Assistant  in  Botany,  Stanford  University,  California. 
WILLIAMS,  FRANCIS  XAVIER;  B.A. 

Curator  of  Insects,  University  of  Kansas,  Lawrence,  Kansas. 
WINSLOW,  MARTHA  MINERVA;  M.A. 

Teacher  of  Physiology  and  Botany,  Pasadena  High  School,  Pasadena,  Cali- 
fornia. 
ZSCHOKKE,  THEODORE  CHRISTIAN;  B.A.,  M.F. 

Forest  Assistant,  United  States  Forest  Service. 


SCIENTIFIC  PAPERS 


THE   VITALITY   OF  THE    SEQUOIA   GIGANTEA  * 
By  WILLIAM  RUSSEL  DUDLEY 

EXISTING  along  the  western  slopes  of  the  Sierra  Nevada  range  in  Cali- 
fornia, in  isolated  groves  from  Placer  County  to  Tulare,  Sequoia 
gigantea  is  a  relict  of  another  age  and  time.  In  these  trees  we  have—- 
with the  Sequoia  sempervirens,  the  redwood  of  the  Coast  Ranges — the 
remnants  of  a  great  genus  that  once  spread  over  all  the  Northern  hemi- 
sphere, as  evidenced  by  fossil  specimens  of  a  considerable  number  of  species 
found  from  France  and  Hungary  to  Spitzbergen,  and  from  Greenland  to 
Oregon  and  Nebraska.  These  fossils  are  found  from  the  Cretaceous,  through 
the  Tertiary,  down  to  recent  times.  How  the  species  have  disappeared  and 
the  individuals  of  the  Big  Trees  have  shrunken  to  the  few  thousands  perched 
among  the  high  salubrious  valleys  of  our  Sierra,  cannot  be  easily  answered. 
The  unknown  complex  of  causes  which  brought  about  the  great  ice  age, 
brought  Sequoia  as  a  race  near  to  extinction;  and  conditions  surrounding  life 
were  so  profoundly  changed,  that  their  former  distribution  could  never  be 
again  restored.  Whatever  the  cause  of  the  present  restriction  of  this  species, 
its  comparative  rarity,  its  inaccessibility,  its  great  size,  its  majesty  and  the 
beauty  if  its  coloring  have  all  served  to  enhance  the  interest  which  we  all 
feel  in  the  California  Big  Tree. 

Indeed  it  is  so  noble,  that  it  has  been  the  subject  of  a  considerable 
amount  of  exaggeration  and  mistaken  comment.  The  statements  that  its 
age  reaches  4,000  to  6,000  years,  and  its  height  exceeds  400  feet  do  not  seem 
to  be  based  on  any  actual  observation.  Nevertheless  it  is  crowned  with 
many  titles  to  greatness,  and  the  most  remarkable  of  all  is  its  relative  ap- 
proach to  immortality.  The  evidence  that  all  living  things  are  finite  is 
so  overwhelming  that  the  mind  is  chastened  with  the  thought  of  it.  But 
the  life  of  a  single  great  tree  of  Sequoia  gigantea,  when  known  clearly,  stirs 
the  imagination  again  to  thoughts  of  what  might  be  attained,  if  disease 
and  the  crushing  weight  of  physical  injury,  as  factors  controlling  life, 
could  be  eliminated.  Certainly  the  oldest  of  the  Big  Trees,  such  as  we 
see  in  the  Calaveras  groves  and  the  forests  of  the  Kings  and  Kaweah  rivers, 
have  the  distinction  of  being  the  oldest,  the  longest  enduring  upon  the  face 
of  the  earth,  of  any  living  organism ;  and  this  is  largely  because  of  their 
freedom  from  disease  and  inherited  weakness  and,  as  I  propose  to  show  a 
little  later,  from  their  marvelous  recuperative  power  in  the  face  of  injury. 


*  Read   by   invitation   before   the   California   Alumni    Association   of    Columbia 
University,  January,   1905. 


34  DUDLEY    MEMORIAL    VOLUME 

The  forests  of  the  Sierra  Nevada  in  October  are  not  dissimilar  in  aspect 
to  those  of  the  Appalachian  mountain  ranges.  Yellowing  oaks  lighten 
the  somber  conifers,  and  crimson  dogwoods  lend  an  aspect  of  brilliancy 
to  the  forest,  unknown  to  the  camper  beneath  its  shade  in  summer.  Even 
Sequoia  exhibits  a  warm  golden  tint,  due  to  a  thousand  small  yellow  branch- 
lets  which  are  maturing  preparatory  to  the  annual  natural  pruning  of  the 
species.  It  was  a  pleasure  to  incidentally  note  these  forest  charms  when 
in  October,  1900,  I  made  my  way  into  the  lumber  camp  belonging  to  the 
Sanger  Lumber  Company  in  the  Converse  Basin  near  the  Kings  River. 
This  mill  is  probably  the  largest  in  capacity  of  any  along  the  forested 
slopes  of  the  Sierra  Nevada.  During  the  previous  month  of  August  it 
cut  200,000  feet  of  lumber  a  day,  or  considerably  above  5,000,000  feet  for 
that  month.  The  records  for  the  other  working  months  of  1900  fell  some- 
what short  of  this  amount,  but  an  enormous  quantity  was  flumed  for  forty 
miles  down  to  the  railroad  at  Sanger,  in  the  San  Joaquin  Valley,  6,000  feet 
below  the  mill.  While  some  of  this  was  pine  and  fir,  the  greater  proportion 
was  made  from  the  giant  trunks  of  the  California  Big  Tree.  Had  a  measur- 
ably large  amount  of  these  trunks  been  utilized  for  lumber  the  cutting  might 
have  been  justified  from  the  lumberman's  point  of  view;  but  frequently 
one-half  to  even  three-fourths  or  seven-eighths  of  the  great  trunks  were 
broken  and  rent  beyond  use  in  falling.  Not  anywhere  in  the  world  is  there 
such  wasteful  lumbering,  and  this  is  a  species  that  above  all  trees,  should 
be  saved  from  the  lumberman! 

The  Converse  Basin,  before  its  deforestation — for  its  forests  have  now 
been  entirely  leveled — presented  for  observation  and  study  the  best  develop- 
ment of  this  rare  coniferous  species  that  existed.  The  trees  were  large  and 
continuous  in  area,  and  this  high  mountain  "basin,"  like  all  others  on  the 
slopes  of  the  Sierras  containing  Sequoias,  is  watered  by  small  brooks  of 
sparkling  spring  water.  Here,  too,  the  streams  soon  plunge  by  cataracts 
into  the  profound  gorge  of  the  Kings  River,  thus  ensuring  excellent  drain- 
age and  good  conditions  for  growth;  and  here  a  brief  visit  in  the  summer 
before  had  shown  me  the  great  number  of  cut  trees  with  logs  and  stumps 
remaining,  which  gave  an  unrivaled  opportunity  to  continue  certain  ob- 
servations already  begun.  My  object,  while  determining  the  age  of  the 
trees  by  means  of  the  number  of  their  annular  layers  of  wood,  was  to 
observe  their  record  year  by  year,  century  by  century,  of  their  behavior  toward 
nutrition,  injury  or  disease. 

The  age  of  a  tree  can  only  be  told  by  counting  the  concentric  rings 
of  growth  on  the  cross-section  of  the  felled  trunk.  The  question  may 
be  asked:  Does  each  ring  represent  a  year's  growth?  A  considerable 


VITALITY    OF    THE    SEQUOIA    GIGANTEA  — ^  DUDLEY  35 

number  of  observations  on  several  species  of  conifers  and  oaks  enables  us 
to  answer  that  it  does,  approximately,  in  those  observed  on  the  Pacific  Coast. 
If  exceptional  seasons  cause  variations  from  this  rule,  the  variations  would 
be  small  in  number  and  not  greatly  affect  the  totals.  During  my  examina- 
tion of  the  felled  trees  of  the  Kings  River,  it  was  a  part  of  my  task  to 
carefully  traverse  these  records  of  growth;  but  I  will  here  give  you  briefly 
only  the  results.  Of  the  various  trunks  of  Sequoia  gigantea  examined  rang- 
ing from  900  years  upward,  the  oldest  possessed  2,425  rings,  or  had  begun 
its  existence  525  years  before  the  Christian  era.  Extended  scrutiny  un- 
doubtedly would  bring  to  light  trees  even  older  than  this,  but  I  do  not 
expect  any  to  exceed  3,000  years  of  age. 

It  has  often  been  inferred  that  the  size  of  a  Big  Tree  bears  an  approxi- 
mately exact  relation  to  its  age.  If  a  tree  exists  eighty  feet  in  circumference 
five  feet  above  the  base,  it  was  inferred,  it  would  be  twice  as  old  as  one 
forty  feet  in  circumference.  This  was  found  to  be  very  far  from  true.  The 
favorite  situation  of  the  larger  trees  is  near  some  hollow,  where  a  tiny 
perennial  spring  brook  is  always  flowing.  The  soil  should  be  good  and 
deep,  but  with  a  large  amount  of  mineral  matter  in  it;  and  above  all,  I 
think  well  drained,  though  always  moist. 

One  tree  occupying  such  a  situation  and  at  the  confluence  of  two  small 
Sierra  Brooks,  was  over  eighty  feet  in  circumference  ten  feet  from  the 
ground,  but  was  only  1,510  years  old,  all  the  rings  being  measurably  thick 
and  uniform.  It  felt  the  effects  neither  of  drouth  nor  of  unusual  precipita- 
tion, and  it  had  never  been  burned  beneath  its  bark. 

On  the  other  hand,  the  tree  which  a  little  later  I  shall  use  as  the  chief 
illustration  of  this  paper,  was  a  small  tree  for  one  of  its  age.  It  stood 
on  a  hillside  not  near  a  stream;  the  influence  of  years  of  abundant  rains 
and  nutrition  were  shown  by  rings  of  fair  degree  of  thickness;  the  effects 
of  years  of  scarcity  were  seen  in  rings  so  thin  that  fifty  of  them  would 
not  cover  an  inch  of  the  tree's  radius.  Moreover,  from  its  unprotected 
situation  it  had  been  seriously  attacked  by  forest  fires,  each  burning  away 
portions  of  its  sap-wood  and  thus  assailing  the  vitality  of  the  plant.  This 
tree  was  only  thirty-nine  feet  in  circumference  ten  feet  from  the  ground, 
but  had  attained  the  age  of  2,171  years  and  a  height  approaching  300  feet, 
although  injury  and  failing  strength  had  resulted  in  a  dead  and  broken 
top  and  reduced  the  tree  to  270  feet  at  the  time  of  its  destruction  in  1900. 

Observations  of  the  greatest  interest,  however,  concerned  the  Big  Tree's 
behavior  toward  severe  injury;  evidences  of  a  remarkable  recuperative  power 
being  found  after  examination  of  the  Sequoias  of  the  Converse  Basin.  The 
effects  of  certain  tremendous  forest  fires  were  registered  in  the  trunks  of 


36  DUDLEY    MEMORIAL   VOLUME 

these  trees,  but  the  record  was  completely  concealed  by  subsequent  healthy 
growth. 

Among  a  number  of  similar  cases  the  most  instructive  record  of  these 
ancient  fires  was  observed  in  the  tree  of  moderate  size — the  one  of  2,171 
years  of  age  above  mentioned.  This  tree,  when  felled,  had  an  enormous 
surface  burn  on  one  side,  occupying  eighteen  feet  of  the  circumference 
and  with  a  height  estimated  at  thirty  feet.  The  fire  had  eaten  through  the 
sap-wood  and  deeply  into  the  heart.  It  was  an  immense  black  scar  and 
an  apparently  irreparable  wound  upon  a  tree  already  advanced  in  age, 
even  for  a  Sequoia.  Yet  this  was  not  the  only  similar  injury  it  had  suffered, 
and  before  we  describe  the  remarkable  life  of  this  tree  as  registered  within, 
let  us  see  how  a  Sequoia  goes  about  the  repair  of  such  appalling  wounds. 

A  burn  on  the  bole  of  a  giant  Sequoia  occurs  usually  on  one  side  only- — 
the  side  toward  the  forest  fire.  It  may  be  a  foot  wide  or  even,  as  in  this 
case,  occupying  an  enormous  area  of  the  trunk  surface;  it  may  be  of 
great  height,  it  may  decrease  the  tree's  vitality  and  yet  not  fatally  injure 
the  individual  thus  attacked.  While  there  is  life  there  is  growth.  After 
the  wound  comes  the  healing;  and  there  is  nothing  more  insistent  (if  we 
may  use  the  word)  in  the  processes  of  plant  life  than  the  attempt  of  a 
strong  tree  to  cover  a  wound  on  its  surface. 

We  have  two  words  in  our  language  in  which  the  pronunciation  and 
even  the  correct  orthography  is  the  same,  but  each  has  a  different  meaning, 
together  with  a  very  distinct,  ancient  and  highly  respectable  ancestry;  I 
mean  the  word  heal.  It  is  a  curious  fact,  moreover,  that  one  of  these  words 
is  properly  applied  to  the  process  of  healing  seen  in  animal  life,  the 
other  describes  the  process  of  covering  a  wound,  such  as  that  adopted  by 
the  tree.  The  first .  and  frequently  used  word  heal  means  to  make  sound, 
and  implies  that,  after  a  wound,  new  tissue  organically  connected  with  the 
old  has  been  formed,  that  the  muscular,  the  circulatory  and  the  nervous 
systems  have  been  extended  in  a  normal  way  from  the  old  to  the  new, 
and  conditions  in  the  once  injured  part  have  been  restored  to  complete 
and  harmonious  working  order.  The  latter  word  heal  means  to  cover  or 
conceal.  It  is  chiefly  observed  in  the  gardener's  art  in  the  expression  "to 
heal  in"  (less  correctly  "heel")  as  applied  to  nursery  seedlings.  It  is  a 
much  rarer  word  than  the  former,  but  good  old  English,  and  we  are  told 
that  it  may  be  traced  back  through  the  German,  the  Gothic  and  even  the 
Latin.  We  remark,  by  the  way,  that  so  far  as  we  know,  not  only  the 
English  word,  but  its  whole  family-tree  clear  back  to  the  Latin  root,  is  no 
older  than  single  individuals  of  Sequoia  gigantea  to-day  standing  in  the 
full  vigor  of  life  in  the  groves  of  Calaveras  and  Kaweah. 


VITALITY    OF    THE  'SEQUOIA    GIGANTEA DUDLEY  37 

It  is  the  latter  word  we  use  in  this  paper.  The  effort  of  an  injured 
tree  is  not  one  to  re-establish  organic  connection  of  the  new  tissue  with  the 
old,  injured  surface  below,  but  wholly  one  to  complete  and  re-establish,  by 
extension,  the  broken  circle  of  growth — the  broken  annual  rings — to  round 
out  the  tree  again  to  its  full  circumference,  to  establish  roots  below,  sup- 
porting and  sap-conducting  tissue  above.  Fortunately  for  the  tree,  it  has 
no  nervous  system  connected  with  delicately  organized  "nerve  centers" ;  no 
circulatory  system  extending  to  every  point  of  its  surface  and  connected  with 
and  controlled  by  a  small  uncertain  organ  deep  within  the  body.  The 
heart-wood  of  a  Big  Tree  is  imperishable  while  the  tree  stands  and  long 
after  it  falls,  unless  attacked  by  fire.  In  the  words  of  the  foreman  of  the 
logging  camp,  "nothing  hurts  the  heart  of  a  redwood — nothing;  it's  always 
sound."  Moreover,  it  is  completely  independent  of  the  living  zones  outside 
of  it,  although  joined  cell  by  cell  with  the  living  tissue.  Its  cells  have 
ceased  to  grow  or  change,  and  the  living  juices  of  the  plant  have  ceased 
to  flow  in  them.  From  the  point  of  view  of  life,  whatever  tissue  in  a  tree 
has  ceased  to  grow  in  every  sense,  has  ceased  to  be  vitally  useful.  Only 
that  tissue  which  is  in  the  process  of  building  is  living,  is  a  vital  part  of 
the  great  organism,  and  this  life  must  exist  in  a  complete  cylinder  forming 
the  outer  tissues  of  the  trunk;  a  circle  constantly,  and  in  the  case  of  a 
Big  Tree,  indefinitely  widening.  If  the  circle  is  broken,  apparently  all  the 
energies  of  life  and  growth  are  directed  toward  closing  it  again,  not  toward 
any  useless  vivification  of  the  dead  cells  of  the  wound  below.  The  burned 
surface  is  dead  tissue,  not  differing  essentially  from  that  of  the  tree's  heart- 
wood,  and  the  healing  of  the  latter  is  only  incidental  to  the  tree's  supreme 
effort  at  the  extension  of  its  living  tissue  over  the  wound  in  order  to  re- 
unite the  margins  of  the  zone  that  should  have  remained  inviolate  and  un- 
broken. 

The  increase  of  the  tree  is  rhythmical,  as  we  have  seen,  accompanying 
the  sun  and  the  seasons.  The  Spring  after  a  wound  has  occurred,  the  tree 
begins  its  effort  toward  healing  by  the  formation  of  a  layer  of  wood  and 
bark  along  all  margins  of  the  burned  area.  This  is  repeated  the  second 
year,  a  layer  of  new  wood  tissue  being  superimposed  upon  that  of  the 
previous  year  along  the  burned  margins.  These  layers  next  the  wound  are 
much  thicker  than  the  ordinary  ones,  the  ring  for  the  same  year  on  the 
side  of  the  tree  opposite  the  burn  being  often  correspondingly  thinner  and 
more  attenuated.  This  process  continues  with  each  returning  season,  and 
the  new  tissue  reaching  inwards  from  all  margins  of  the  injury  takes  on 
the  form  of  solid  folds  of  wood  growing  uniformly  broader  on  each  side, 
the  black  char  narrower,  year  by  year.  There  is  no  organic  union,  however, 


38  DUDLEY    MEMORIAL   VOLUME 

between  the  new  wood  of  the  folds  and  the  wood  of  the  charred  surface 
underneath  them,  no  healing  at  this  point  of  contact,  in  the  ordinary  sense 
of  the  word ;  but  there  is  effectual  covering,  or  healing  in  the  rarer  sense, 
according  to  the  tree  trunk's  way.  Sometimes,  from  the  attack  of  insects 
on  the  rapidly  formed  wood  of  the  folds,  these  folds  die.  There  is  no 
surgeon  present  to  cut  away  this  dead  tissue,  but  the  tree  patiently  begins 
to  form  a  new  fold  to  cover  the  dead  one.  In  a  species  with  the  ordinary 
span  of  life  the  delay,  this  waste  of  effort,  might  be  fatal  to  the  final 
closing  of  the  wound.  Not  so  with  the  Big  Tree,  to  whom  a  score  of  years 
is  as  one.  The  first  fold  is  overtaken  and  passed — in  one  case  it  took  just 
fifty  years  to  do  it — and  sooner  or  later  the  two  folds  from  opposite  sides 
touch  one  another ;  a  few  years  more  and  the  bark  is  pinched  out,  the  charred 
surface  is  entirely  covered,  and  finally  the  annular  layers  become  continuous 
around  the  entire  circumference,  each  resuming  a  normal  thickness  through- 
out. The  process,  which  has  drawn  on  all  the  resources  of  the  plant,  it 
may  be  for  scores  of  years,  it  may  be  for  centuries,  is  completed.  The 
wound  is  healed!  This  is  a  momentous  event,  yet  only  the  spectacle  of  a 
perfected  cylinder  with  the  splendid  circumference  of  forty  or  sixty  or 
ninety  feet,  of  living  tissue  through  which  the  sap  of  the  tree  can  pass,  to 
a  considerable  extent  laterally  as  well  as  vertically,  is  the  result;  only  con- 
tinuous healthful  growth  and  unbroken  increase,  the  most  inspiring  of  all 
spectacles. 

In  the  life  history  of  a  Big  Tree  such  injury,  such  prolonged  but  com- 
plete and  thorough  repair  may  occur  not  only  once  but  several  times,  and 
yet  all  evidence  of  the  various  catastrophies  be  entirely  obliterated  except 
for  the  thin  cavities,  each  with  one  charred  surface,  and  the  peculiar  struc- 
ture of  the  repair  layers  deep  within  the  undecayed  heart  of  the  tree.  When 
the  tree  of  2,171  years  of  age  was  cut,  in  addition  to  the  great  burn  on  its 
trunk  eighteen  feet  in  width,  the  record  of  three  other  fires  was  revealed. 
The  history  of  the  tree  was  as  follows: 

It  began  its  existence  271  B.  C. 

At  the  beginning  of  the  Christian  era  it  was  estimated  to  be  already 
about  twelve  feet  in  circumference  just  above  the  base. 

At  516  years  of  age  (A.  D.  245)  occurred  a  burning  three  feet  in 
width  on  the  trunk. 

One  hundred  and  five  years  were  occupied  in  healing  this  wound. 

One  thousand,  one  hundred  and  ninety-six  years  without  injury  followed. 

At  1,712  years  of  age  (A.  D.  1441)  occurred  a  second  burning,  making 
two  wounds  of  one  and  two  feet  each  in  width.  Each  had  its  own  system 
of  healing. 


VITALITY    OF    THE    SEQUOIA    GIGANTEA DUDLEY  39 

One  hundred  and  thirty-nine  years  of  growth  followed,  including  the 
time  occupied  by  the  covering  of  the  two  wounds. 

At  1,851  years  of  age  (A.  D.  1580)  occurred  another  fire,  causing  a 
burn  on  the  trunk  two  feet  wide,  which  took  fifty-six  years  to  heal. 

Two  hundred  and  seventeen  years  of  growth  followed  the  fire. 

When  the  tree  was  2,068  years  old  (in  1797)  a  tremendously  aggressive 
fire  attacked  the  trunk  (perhaps  aided  by  the  burning  stem  of  a  neighbor- 
ing pine  or  fir)  and  burned  the  great  scar  eighteen  feet  wide  with  a  height 
estimated  at  thirty  feet.  The  103  years  which  had  elapsed  since  1797  had 
reduced  this  to  fourteen  feet  in  width.  If  the  same  rate  of  growth  con- 
tinued without  interruption — a  hazardous  estimate — and  the  tree  had  been 
in  possession  of  the  United  States  and  under  its  protection,  the  wound  might 
have  been  closed  in  three  and  one-half  centuries  more,  or  about  the  year  2250. 
Four  centuries  and  a  half  to  repair  in  one  tree  the  results  of  one  forest 
fire!  If  the  tree  had  been  a  younger  tree,  less  the  victim  of  previous 
fires,  we  are  convinced  that  .such  a  healing  would  be  possible.  In  any  case 
Sequoia  gigantea  practically  stands  alone,  sublime  among  living  objects  in 
its  ability  to  withstand  an  injury  of  this  magnitude,  and  to  endure  a  sufficient 
length  of  time  for  its  complete  recovery. 

It  is  to  be  noted  that  in  the  trunk  next  to  each  of  the  three  older 
burns  described,  there  is  a  thin  cavity  chiefly  occupied  by  the  charcoal  of 
the  burned  surface  (some  of  that  formed  in  245  was  brought  away),  and 
that  this  produced  a  pathologically  protective  covering,  no  doubt  calculated 
as  well  as  any  to  prevent  decay  during  the  long  period  consumed  in  cover- 
ing the  wound  with  healthy  tissue.  But  this  will  not  account  for  all  this 
superb  resistance  to  the  attack  of  insect,  fungus,  ferment  or  microbe.  Burned 
areas  of  other  trees  have  the  same  charred  surface,  but  no  oak  or  sycamore, 
pine  or  Douglas  spruce  under  similar  conditions  would  remain  so  long  with- 
out being  attacked  in  this  region  by  some  cause  of  decay.  There  is  some- 
thing in  the  sap  of  the  Big  Tree  that  is  an  elixir  of  life,  something  deposited 
in  the  lignified  cells  of  the  normally  formed  layers  of  wood  that  resists  in 
an  unexampled  way  the  dreaded  "tooth  of  time."  The  wound  is  finally 
covered — not  healed,  in  the  surgeon's  sense — the  new  tissue  formed  above 
it  is  thickened,  the  tree  is  rounded  to  its  original  fullness,  bark  and  wood 
become  continuous  about  the  whole  circumference,  the  latter  forming  in 
rings  of  normal  uniformity,  the  old  healthful  symmetry  of  life  is  re-estab- 
lished, and  no  outward  sign  of  distortion  exists,  or  even  a  scar  from  the  old 
injury.  Nevertheless,  well  within,  and  as  the  centuries  pass,  deeper  and 
deeper  within  the  heart  of  the  tree  the  wound  exists  unchanged  and  there- 
fore no  source  of  decay. 


40  DUDLEY    MEMORIAL    VOLUME 

Again,  it  is  to  be  observed  from  the  notes  of  the  tree's  yearly  growth, 
that  after  it  was  out  of  its  first  youth,  the  periods  of  most  vigorous  increase 
were  after  the  successive  burnings  and  during  the  periods  of  healing.  In 
part  this  accounted  for  the  lessened  area  of  the  tree's  live  circumference ; 
but  as  this  thickening  of  the  rings  appears  to  continue  after  complete  healing 
has  taken  place,  when  the  tree  is  again  forming  tissue  over  its  entire  cir- 
cumference, and  as  this  phenomenon  was  seen  in  other  trees  similarly  in- 
jured, one  is  led  to  believe  that  an  increased  activity  in  the  tree's  life  had 
been  occasioned  not  by  the  burn,  but  from  the  effort  at  healing  and  recover- 
ing from  what  threatened  to  be  a  vital  wound;  and  that  this  activity  led 
to  more  vigorous  growth.  It.  is  a  curious  fact,  moreover,  that  in  the  middle 
of  the  long  period  of  freedom  from  fire,  from  245  to  1441,  a  period  of  nearly 
twelve  centuries,  the  tree  made  its  least  relative  increase  in  diameter.  Peace 
and  apparent  prosperity  had  been  coincident  with  a  sluggish  growth  if  they 
had  not  been  the  cause  of  it. 

About  three  and  one-half  inches  was  the  most  frequent  amount  of  radial 
growth  during  one  century,  a  total  increase  of  about  seven  inches  in  the 
diameter  of  the  stem;  but  during  the  first  six  hundred  years  its  average  was 
five  inches  (a  total  increase  of  ten  inches  in  diameter  each  century)  and 
during  the  first  and  fourth  centuries  of  this  tree's  existence  its  radial  increase 
was  six  inches  in  each  case.  During  the  seventh,  eighth,  ninth,  tenth  and 
eleventh  centuries  the  increase  in  the  radius  of  the  stem  was  between  two 
and  three  inches  only  per  century.  It  was  during  this  period,  from  the 
seventh  to  the  twelfth  century  of  its  existence — the  period  of  greatest  depres- 
sion in  the  apparent  vitality  of  the  tree — that  the  rings  or  annular  layers  be- 
came so  thin  that  it  was  impossible  to  count  them  without  the  aid  of  a  lens : 
over  forty  layers  were  frequently  found  in  one  inch  of  radial  line,  and  in 
two  cases  apparently  there  were  fifty-two  and  fifty-four  layers  in  each  inch. 

I  cannot  help  thinking  we  are  here  in  the  presence  of  one  of  the  most 
remarkable  products  of  the  globe,  not  excepting  those  of  human  civilization. 
Almost  no  structure  erected  by  human  hands  has  come  down  to  us  intact 
through  the  lifetime  of  a  Sequoia;  and  the  few  we  can  admire  are  hewn 
from  inanimate  marble  or  granite  and  cannot  be  compared  to  a  living  or- 
ganism, vast  in  life  and  complete  in  the  records  of  every  year  of  its  existence. 
An  empire  or  republic  may  be  compared  to  the  life  of  these  great  trees. 
But  what  empire  or  republic  has  lived  for  twenty-five  centuries?  None 
worthy  of  the  name,  and  certainly  none  among  those  of  the  Aryan  civilization. 
Then  in  the  building  of  a  Sequoia,  no  blood  has  been  shed  through  all 
its  twenty-five  hundred  years,  no  injustice  or  oppression  has  secured  the 
means  necessary  for  its  construction,  no  hate  or  strife  has  been  engendered, 


VITALITY    OF    THE    SEQUOIA    GIGANTEA DUDLEY  41 

no  accident  occasioning  pain  or  suffering  or  the  extinction  of  human  life 
has  left  a  stain  on  the  history  of  its  growth.  Tragedies  and  great  passions, 
as  we  have  seen,  have  crossed  its  silent  life,  but  they  have  been  the  elemental 
passions  of  fire  and  storm,  clean  and  wholesome,  and  the  tree  has  been 
stimulated  by  them  to  a  greater  and  more  vigorous  growth.  Indeed,  there 
is  something  sublime  in  the  patience  of  the  task  and  the  completeness  of  its 
execution  when,  after  centuries  of  slow  rebuilding,  we  see  every  outward  trace 
of  its  injuries  eliminated  and  a  robust  and  uninterrupted  life  again  at- 
tained. 

Mr.  James  Bryce,  in  his  'sketch  of  the  life  of  the  late  Lord  Acton, 
Professor  of  History  at  the  University  of  Cambridge,  says:  "Twenty  years 
ago,  at  midnight  in  his  library  at  Cannes  he  expounded  to  me  how  a  history 
of  Liberty  might  be  written  and  in  what  way  it  might  be  made  the  central 
thread  of  all  history.  He  spoke  like  a  man  inspired,  seeming  as  if  from  some 
mountain  summit  high  in  air  he  saw  beneath  him  the  far  winding  path  of 
human  progress,  from  dim  Cimmerian  shores  of  prehistoric  shadow,  into  the 
fuller,  yet  broken  and  fitful  light  of  the  modern  time.  *  *  *  It  was 
as  if  the  whole  landscape  of  history  had  been  suddenly  lit  up  by  a  burst  of 
sunlight.  I  had  never  heard  from  any  other  lips  any  discourse  like  this,  nor 
from  his  did  I  hear  the  like  again." 

The  impression  made  was  not  dissimilar,  on  that  cloudless  October 
afternoon  with  the  crimson  leaves  of  the  dogwood  and  the  yellow  oak  fall- 
ing silently  in  the  Sierra  forests,  as  one  patiently  wrought  out  with  lens, 
measure,  pencil  and  camera  the  great  history  of  the  Sequoia  above  named, 
year  by  year,  century  by  century;  centuries  of  peace,  years  of  tragedy,  and 
again  centuries  of  stimulated  growth.  It  was  as  if  the  whole  landscape 
of  life,  from  the  dim  prehistoric  forests  until  now,  "had  been  suddenly  lit 
up  by  a  burst  of  sunlight."  I  had  never  heard  from  any  other  lips  any  dis- 
course like  this,  nor  from  this  fallen  seer  and  patriarch  could  I  hear  the  like 
again. 

During  the  past  ten  years  hardly  a  season  has  passed  but  I  have  camped 
among  the  Sequoias.  I  am  glad  to  say  I  have  visited  nearly  all  the  groves, 
but  I  regret  to  say  that  a  considerable  proportion  of  them  is  in  private 
hands ;  some  have  been  leveled  already  and  the  mills  are  busy  in  not  less 
than  four  others,  notwithstanding  there  is  little  profit  in  the  lumbering. 
These  groves  of  Sequoias  form  a  question  apart  from  the  ordinary  questions 
of  forestry.  In  the  heart  of  the  Sierra  forests  they  are,  it  is  true,  an  im- 
portant part  of  the  protective  forest  cover  of  the  headwaters  of  California 
rivers;  but  I  believe  you  will  now  join  me  in  the  assertion  that  they  have 
an  interest  for  the  citizens  of  California,  for  the  cultivated  traveler  and  the 


42  DUDLEY    MEMORIAL   VOLUME 

scientific  man,  far  beyond  that  of  the  other  trees  of  the  forest.  The  United 
States  should  own  and  properly  protect  every  one  of  them.  Senator  Hoar, 
of  Massachusetts,  once  said  if  the  Calaveras  Groves  were  in  Massachusetts, 
she  would  herself  buy  them  and  not  ask  the  National  government  to  pur- 
chase them.  There  are  some  things,  however,  that  are  the  natural  heritage 
of  a  nation,  and  the  Sequoia  gigantea  is  one  of  them.  We  would  rather 
see  the  Yellowstone  National  Park  and  the  Grand  Canyon  of  the  Colorado 
under  the  protection  of  the  United  States  than  that  of  any  state,  not  ex- 
cepting California  or  Massachusetts.  If  the  Sequoias  are  among  the  most 
remarkable  objects  on  the  globe,  if  they  are  the  best  calculated,  as  we  can 
show,  of  any  living  organism  to  throw  light  on  certain  problems  of  scientific 
inquiry,  then  a  nation  should  own  them  and  their  preservation  should  be 
a  matter  of  national  pride. 

We  make  every  effort  to  preserve  the  manuscript  of  our  great  Anglo- 
Saxon  and  American  charters;  nevertheless  the  ink  fades,  the  parchment 
crumbles  and  they  disappear,  except  from  the  lives  of  just  men.  We  house 
the  archives  of  our  wars  in  buildings  of  great  cost,  maintained  with  great 
care,  yet  all  these  are  on  paper  that  is  more  perishable  than  the  parchment 
of  our  charters.  In  these  great  trees,  however,  we  have,  deep  in  their  annual 
rings,  records  which  extend  far  beyond  the  beginnings  of  Anglo-Saxon 
peoples,  beyond  even  the  earliest  struggles  for  liberty  and  democracy  among 
the  Greeks,  the  first  of  the  Indo- Europeans  to  crystallize  into  national  life 
through  the  pressure  of  this  struggle.  The  records  are  those  of  forest 
conflagrations,  of  the  vicissitudes  of  seasons,  of  periods  of  drouth  and  periods 
of  abundant  and  favoring  rains,  and  we  might  find  next  to  the  charcoal  of 
some  trunk  scar,  centuries  old,  the  stone  inplements  belonging  to  the  ancient 
aboriginal  inhabitants  of  Western  America.  Practically  none  of  these  records 
have  yet  been  studied.  Let  the  nation  purchase  these  trees  of  the  Cala- 
veras groves — among  the  largest  of  all  those  still  remaining  alive — let  it 
take  them  as  a  right  and  a  duty,  not  parleying  with  the  cupidity  of  an 
owner  who  has  done  nothing  to  increase  their  value;  let  it  gradually  gather 
under  its  protection  all  the  groves  of  Sequoia,  now  in  alien  hands,  and  care 
for  them  all  intelligently.  When  the  oldest  of  trees  succumb  and  die,  as 
from  past  injuries  they  must  do  in  time,  then  let  them  be  felled;  and  in- 
stead of  being  sold  or  burned  with  criminal  indifference  to  their  real  value, 
as  at  the  present  day,  may  their  records  be  read  and  recorded  by  skilled 
hands  and  interpreted  by  the  best  intelligence ;  and  finally  may  their 
massive  timbers,  of  wonderful  fineness,  uniformity,  luster,  color  and  beauty, 
be  used  only  in  the  interior  of  a  nation's  buildings,  in  places  which  shall 
the  longest  endure. 


THE  MORPHOLOGY  AND  SYSTEMATIC  POSITION  OF 
CALYCULARIA    RADICULOSA    (Steph.) 

By  DOUGLAS  HOUGHTON  CAMPBELL,  Professor  of  Botany 

THE  classification  of  the  so-called  anacrogynous  Jungermanniales,  an 
important  group  of  liverworts,  is  at  present  in  a  very  unsatisfactory 
condition,  and  much  remains  to  be  done  before  the  true  relationships 
of  the  members  of  this  group  can  be  satisfactorily  settled.  A  recent  attempt 
has  been  made  toward  a  better  classification  of  the  liverworts  by  Cavers1 
and  this  is  a  distinct  advance  upon  the  classifications  which  have  heretofore 
been  accepted.  There  are,  however,  a  number  of  forms  whose  relationships 
a*re  still  by  no  means  clear,  and  among  these  is  a  rare  liverwort  from  Java 
originally  described  as  Calycularia  radiculosa.  Schiffner2  speaks  of  the 
plant  as  an  extremely  rare  one,  but  during  a  stay  in  Tjibodas  where  the 
plant  had  been  collected  before,  the  writer  succeeded  in  finding  it  a  number 
of  times.  The  plants  did  not  grow  in  large  masses  but  'were  associated  with 
various  other  liverworts  growing  on  the  trunks  of  trees.  The  general  aspect 
of  the  plant  (see  Figs.  1  and  2)  is  very  much  like  that  of  the  creeping 
forms  of  the  genus  Blyttia.  Pallamcinia  (Bly.ttia)  Levieri,  a  common  species 


Fig.  1.  Three  male  plants  of  Calycularia 
radiculosa.  Steph.  x  3.  $ ,  antheridial  recep- 
tacle. 


1  The  inter-relationships  of  the  Bryophyta.     New  Phytologist  Reprint,  No.  4, 


1911. 


2  Die  Hepaticae  der  Flora  Von  Buitenzorg,  1900. 


44 


DUDLEY    MEMORIAL   VOLUME 


of  the  same  region,  much  resembles  the  plant  in  question,  but  the  latter  is 
readily  distinguishable  on  account  of  the  numerous  dark  reddish  purple 
rhizoids. 


Fig.  2.  A,  female  plant  1x3;  the  thallus  has  begun  active  growth  again  and 
developed  a  second  archegonial  receptacle,  o.1,  in  the  new  portion.  B,  part  of  one 
of  the  involucral  scales,  slightly  magnified.  C,  female  plant  bearing  a  young 
sporophyte,  sp.  D,  plant  with  mature  sporophyte.  E,  open  capsule  with  the  valves 
entirely  separated  at  the  apex.  In,  involucre ;  per,  perianth. 


The  material  collected  by  the  writer  was  sufficient  to  make  possible  a 
pretty  complete  study  of  the  structure  and  development  of  the  plant,  except 
the  earlier  stages  of  the  sporophyte  which  were  wanting  in  the  specimens 
collected. 


CALYCULARIA    RADICULOSA CAMPBELL 


45 


The  genus  Calycularia  as  generally  understood  comprises  four  species 
of  thallose  liverworts  of  rather  unusual  distribution.  One  of  these,  C.  laxa, 
occurs  in  arctic  Siberia,  two,  C.  crispula  and  C.  Birmensis,  are  found  in  India 
and  Burmah,  while  the  other,  C.  radiculosa,  occurs  in  Java.  The  question  has 
been  raised  whether  the  latter  species  really  should  be  united  with  the 
others.  Schiffner3  after  an  examination  of  the  plant,  concluded  that  it 
should  be  placed  in  the  genus  Morkia,  a  genus  sometimes  regarded  as  a 
section  simply  of  the  larger  genus  Pallavicinia.  The  material  upon  which 
the  present  account  is  based  was  collected  by  the  writer  in  Java  in  1906 
while  staying  at  Tjibodas,  one  of  the  stations  where  the  plant  had  originally 
been  collected. 


GENERAL    MORPHOLOGY 

The  plants  are  dioecious,  the  male  plants  ( Fig.  1 )  being  decidedly  smaller 
than  the  females  (Fig.  2).  Antheridia  may  be  developed  while  the  plants 
are  not  more  than  5  mm.  in  length,  but  the  male  plants  may  reach  a  length 
of  12-15  mm.  The  female  plants  are  two  or  three  times  as  long  as  the  males 
and  may  reach  a  length  of  about  30  mm.,  with  a  breadth  of 'about  14  mm. 


Fig.  3.  A,  cross-section  of  the  thallus,  showing  the  thickened  mid-rib  and 
rhizoids,  r,  x  about  30.  B,  cells  from  the  ventral  side  of  the  thallus,  showing  the 
mycorrhiza,  m,  extending  from  the  rhizoid,  r,  into  the  inner  cells  of  the  thallus, 
x  about  400.  C,  an  inner  cell  of  the  thallus  invaded  by  the  mycorrhiza ;  the  nucleus 
of  the  cell  is  still  intact.  D,  oogonium-like  enlargement  of  a  mycorrhizal  filament. 


3  Osterreichische  Botanische  Zeitschrift,  Feb.,   1901. 


46 


DUDLEY    MEMORIAL   VOLUME 


They  are  usually  not  branched  but  may  be  forked  once  (Fig.  1,  C).  There 
is  a  deep  sinus  in  front  within  which  lies  the  growing  point  of  the  thallus. 
A  thick  midrib  is  developed  strongly,  projecting  on  the.  lower  side  where 
its  ventral  surface  is  covered  with  numerous  deep  purple-red  rhizoids.  The 
margin  of  the  thallus  is  more  or  less  strongly  undulate  and  folded,  but 
these  undulations  are  hardly  distinct  enough  to  be  called  leaves.  The  whole 
aspect  pf  the  plant  is  very  much  like  certain  species  of  Pallavicinia,  and  also 
suggests  the  Japanese  genus  Makinoa.4  A  section  of  the  thallus  (Fig.  3,  A) 
shows  that  the  midrib  comprises  about  a  dozen  cells  in  thickness,  but  there 
is  no  trace  of  the  conducting  strands  of  tissue  which  are  a  constant  character 
in  Pallavicinia.  In  Eupallavicinia  (Blyttia)  there  is  a  single  very  con- 
spicuous axial  strand,  while  in  Morkia,  according  to  Cavers,  there  are  de- 
veloped two  strands  which  are  however  much  less  strongly  developed  than 
in  Blyttia.  In  the  character  of  the  midrib,  therefore,  Morkia  seems  to  be 
somewhat  intermediate  in  character  between  Calycularia  radiculosa  and 
Blyttia. 

The  wings  of  the  thallus  are  composed  of  a  single  layer  of  cells  in  the 
marginal  region,  but  toward  the  midrib  the  wings  are  composed  of  two  or 


\ 


Fig.  4.  A,  vertical  section  of  the  thallus  apex,  in  which  there  are  dorsal  and 
ventral  segments  cut  off  from  the  apical  cell.  B,  C,  two  consecutive  sections  from 
a  thallus  apex,  in  which  a  single  basal  segment  is  cut  off.  D,  E,  two  nearly 
horizontal  sections,  showing  the  appearance  of  the  apical  cell,  x,  when  seen  from 
above,  x  225,  h,  ventral  glandular  hairs. 


*  Miyake,  K. ;  Makinoa,  A  New  Genus  of  Hepatic*.    Bot.  Mag.,  Vol.  13,  1899. 


CALYCULARIA   RADICULOSA CAMPBELL 


47 


sometimes  even  of  three  layers  of  cells.  In  this  respect  Calycularia  radiculosa 
perhaps  more  nearly  resembles  Makinoa  than  it  does  Pallavicinia. 

A  characteristic  feature  of  the  thallus  is  the  presence  in  the  older  por- 
tions of  an  endophytic  fungus  or  mycorrhiza,  very  much  like  that  found 
in  the  subterranean  prothallia  of  various  pteridophytes.  A  similar  mycor- 
rhiza, however,  has  been  found  also  by  the  writer  in  various  green  fern  pro- 
thallia, and  it  is  also  known  to  occur  in  various  other  liverworts. 

The  fungus  both  in  its  structure  and  manner  of  growth  resembles  more 
closely  the  mycorrhiza  described  by  the  writer5  in  Ophioglossaceae.  As  in 
the  Ophioglossaceae  there  were  occasionally  found  oogonium-like  enlarge- 
ments (Fig.  3,  D)  which  may  have  been  perhaps  special  reproductive  bodies, 
but  this  could  not  be  positively  demonstrated.  The  genus  Calycularia  is 


Fig.  5.    A,  upper  surface  of  a  male  plant,  x,  apex  of  the  thallus.    A^,  under 
surface,  showing  the  antheridia,  $  ,  surrounded  by  the  laciniate   scales,  sc.   x   15. 

B,  median  section  of  the  thallus,  showing  the  apex,  x,  and  the  antheridia,  $  ,  x  40. 

C,  a  horizontal  section  of  the  antheridial  receptacle,  sc,  scales.    D,  scales  showing 
the  laciniate  margins. 


5  Campbell,   D.   H. 
Washington.     1911. 


The   Eusporangiatse.      Pub.    140.      Carnegie   Institution  of 


48  DUDLEY    MEMORIAL   VOLUME 

described  as  having  upon  the  ventral  surface  leaf -like  scales  or  amphigastria, 
but  Schiffner  found  that  these  were  not  present  in  Calycularia  radiculosa, 
and  the  writer's  investigations  confirm  this.  These  leaf-like  scales  are  re- 
placed by  multi-cellular  hairs  (Fig.  4,  Bh),  such  as  are  common  on  many 
other  thallose  Jungermanniales.  The  terminal  cell  is  enlarged  and  probably 
secretes  mucilage  for  the  protection  of  the  thallus  apex. 

The  latter  is  turned  strongly  upward  (Figs.  5,  B;  8,  A)  and  it  is 
almost  impossible  to  make  satisfactory  sections  parallel  with  the  surface 
of  the  thallus.  Figures  4,  D  and  5,  B  show  sections  which  are  approximately 
parallel;  but  as  these  are  somewhat  oblique,  the  apical  cell  appears  some- 
what shorter  than  it  really  is.  In  this  view  it  appears  somewhat  oblong  in 
outline,  and  it  is  evident  that  segments  are  cut  off  both  from  the  lateral  and 
from  the  basal  portions.  In  vertical  horizontal  sections  the  apical  cell  shows 
certain  variations,  resembling  in  this  respect  the  genus  Pellia.  While  Pellia 
epiphylla  has  an  apical  cell  with  a  single  basal  segment  extending  the  whole 
length  of  the  thallus,  in  P.  calycina  a  vertical  section  shows  two  sets  of  seg- 
ments, dorsal  and  ventral,  such  as  occur  in  certain  species  of  Pallavicinia,  as 
well  as  in  the  Marchantiales  and  in  Anthocros.  In  Calycularia  radiculosa 
both  of  these  types  were  found.  The  type  found  in  Pellia  epiphylla  (See 
Fig.  4,  B,  C)  were  common  in  the  smaller  plants,  but  it  was  not  at  all  clear 
whether  there  really  is  any  definite  relation  between  the  thickness  of  the 
thallus  and  the  form  of  the  apical  cell.  The  second  type  is  shown  in 
Figure  4,  A. 

Where  branching  takes  place  it  seems  to  be  a  true  dichotomy,  but  whether 
one  of  the  branches  retains  the  original  apical  cell  or  whether  two  new  apical 
cells  are  developed,  was  not  investigated. 

THE  MALE  PLANT. 

The  male  plants  (Fig.  1)  are  usually  quite  short,  often  being  scarcely 
longer  than  broad,  and  as  we  have  already  stated,  antheridia  are  sometimes 
found  upon  plants  which  are  not  more  than  5  mm.  in  length.  The  antheridia 
are  in  small  groups,  seldom  more  than  ten  together,  and  are  much  less  nu- 
merous than  is  the  case  in  either  Morkia  or  Blyttia.  In  the  restriction  of 
the  antheridia  to  such  a  limited  region  the  plants  suggest  Makinoa,  but  the 
whole  antheridial  group  is  not  subtended  by  a  common  envelope  as  in  Makinoa, 
though  the  antheridia  occupy  a  more  or  less  well-marked  depression  or  cavity 
upon  the  dorsal  surface  of  the  midrib.  Each  antheridium  is  subtended  by  a 
much  laciniated  scale.  The  scales  are  often  more  or  less  confluent,  so  that 
imperfect  chambers  are  formed  (Fig.  5,  C)  about  each  antheridium.  As  a 
rule,  only  one  receptacle  occurs  upon  the  plant,  but  in  a  few  of  the  larger 


CALYCULARIA   RADICULOSA CAMPBELL 


49 


ones,  where  old  groups  of  antheridia  were  present,  a  second  younger  group 
was  occasionally  found  near  the  apex;  and  in  the  rare  instances  where  the 
thallus  forks,  each  branch  may  bear  an  antheridial  receptacle.  (Fig.  1,  C.) 

The  antheridia  (Figs.  5  and  6)  are  short-stalked  nearly  globular  bodies 
and  closely  resemble  those  of  Pallavicinia,  and  as  usual  they  are  formed  in 
acropetal  succession  alternately  right  and  left  of  the  apex  of  the  thallus. 

The  earliest  stages  were  not  found,  .so  that  it  is  impossible  to  say 
whether  the  early  divisions  correspond  to  those  observed  in  other  genera,  but 
as  the  young  antheridia  resemble  so  closely  those  of  Pallavicinia,  it  is  to  be 


Fig.  6.  Development  of  the  Antheridium. 
A,  median  section  of  young  antheridium,  x  about 
225.  B,  cross-section  of  the  antheridium  of  about 
the  same  age.  C,  an  older  antheridium.  D,  E, 
two  nearly  ripe  antheridia,  x  about  90. 


50 


DUDLEY    MEMORIAL    VOLUME 


expected  that  the  early  stages  would  conform  to  the  usual  type  found  in  the 
J  ungermanniales. 

In  his  description  of  the  genus  Calycularia  given  in  Engler  and  Prantl's 
Natiirliche  Pflanzenfamilien,  Schiffner  states  that  the  antheridium  has  a 
single  celled  stalk,  but  in  his  later  description  of  C.  radiculosa,  given  in  his 
work  on  the  Liverworts  of  Buitenzorg  he  says  that  he  did  not  see  the  male 
plants,  so  that  this  description  would  not  apply  to  that  species,  and  as  we 
shall  see,  the  stalk  is  multicellular,  very  much  like  that  of  Pallavicinia. 

Figure  6,  A  shows  a  longitudinal  section  of  the  youngest  perfect 
antheridium  that  was  found.  The  short  stalk  shows  in  sections  two  rows  of 
cells  and  the  upper  portion  shows  a  mass  of  young  spermatogenous  cells 
surrounded  by  a  single  layer  of  sterile  cells.  Figure  6  B  shows  a  cross-section 
of  an  antheridium  of  about  the  same  age.  Sometimes  the  stem  of  the 
antheridium  is  more  slender  and  may  have  a  single  basal  cell  (Fig.  6  E) 
which  often  becomes  very  much  elongated. 

Before  the  final  division  of  the  spermatogenous  cells  they  are  polygonal 
in  outline,  with  dense  contents  usually  more  or  less  contracted,  but  how  far 
this  is  normal  and  how  far  it  is  due  to  artificial  shrinkage  could  not  be 
determined.  The  cell  walls  are  clearly  defined.  The  nucleus  is  conspicu- 
ous and  stains  strongly.  The  contents  appear  somewhat  finely  granular,  the 


Fig.7.  Spermatogenesis.  All  figures  magnified  about  750.  A,  B,  spermatogenic 
cells,  just  before  the  final  mitosis;  in  B  can  be  seen  two  small  bodies,  b,  which  are 
probably  the  young  blepharoplasts.  C,  D,  stages  in  the  development  of  the  sper- 
matozoid.  E,  I,  M,  show  the  pair  of  spermatocytes  seen  from  the  side,  the  others 
are  mostly  single  spermatocytes  seen  from  above;  b,  blepharoplast ;  c,  cilia;  n, 
nucleus  of  the  young  spermatozoid.  Figs.  L,  M,  show  the  mature  spermatozoids. 


CALYCULARIA   RADICULOSA CAMPBELL  51 

nuclear  reticulum  not  being  clearly  visible,  and  there  are  a  number  of  bodies, 
one  of  which  is  probably  the  nucleus,  although  it  is  possible  that  there  may 
be  more  than  one  nucleus  (Figs.  7  A,  B).  In  a  number  of  cases  shortly 
before  the  division  to  form  the  spermatocytes,  there  could  be  seen  two  minute 
granules  (Figs.  7  B,  b),  sometimes  surrounded  by  a  colorless  area  and 
perhaps  representing  the  young  blepharoplasts,  but  the  small  size  of  these, 
and  the  presence  of  other  granular  bodies  in  the  cytoplasm  makes  one  hesitate 
to  assert  positively  that  these  really  were  blepharoplasts,  especially  as  no 
division  stages  were  found,  and  the  relation  of  these  bodies  to  the  nuclear 
spindle  could  not  be  determined. 

The  further  development  of  the  spermatozoids,  which  are  unusually  large 
in  this  species  and  therefore  favorable  for  study,  is  on  the  whole  much  like 
that  of  Pellia.  After  the  final  nuclear  division  a  definite  cell  wall  is 
formed  between  the  spermatozoids,  the  division  not  running  diagonally,  as 
described  by  Ikeno  for  Marchantia,  but  dividing  the  cell  into  two  approxi- 
mately hemispherical  ones.  (Fig.  7  E,  I.) 

In  the  youngest  stages  met  with  (Figs.  7  C,  D)  the  nucleus,  which  now 
appears  somewhat  coarsely  granular,  was  still  unchanged  in  form.  The  cyto- 
plasm was  often  contracted  but  not  always  so,  and  in  the  cytoplasm,  some- 
times at  the  periphery,  but  quite  as  often  near  the  nucleus,  could  be  seen  the 
blepharoplast  (Fig.  7,  C,  D,  b),  which  was  already  extended  into  a  delicate 
band.  No  cases  were  found  where  the  blepharoplast  showed  its  original 
round  form.  A  careful  examination  of  the  nucleus  at  this  time  shows  that 
it  is  decidedly  flattened  in  the  plane  of  the  division  wall  between  the  two 
spermatocytes,  so  that  it  appears  oval  when  seen  from  the  side  but  circular 
when  viewed  from  above. 

No  signs  of  any  body  equivalent  to  the  so-called  "Nebenkorper,"  or  the 
"accessory  body"  described  by  Wilson6  for  Pellia  could  be  seen,  and  such  a 
body  is  probably  quite  wanting  in  Calycularia  radiculosa.  In  a  slightly  later 
stage,  however  (Figs.  E,  F,  G),  there  could  often  be  seen  what  appeared 
like  cytoplasm  extending  beyond  the  nucleus  and  connecting  with  the  ble- 
pharoplast. Strassburger  refers  to  such  a  structure  in  his  somewhat  brief 
account  of  Pellia. 

The  nucleus  now  begins  to  elongate  and  to  increase  notably  in  size,  hav- 
ing the  form  of  an  almost  homogenous  crescent-shaped  body  when  seen  from 
the  side.  The  forward  end  of  the  crescent  is  somewhat  more  prominent 
than  the  posterior  end  and  extends  into  the  cytoplasmic  prominence,  con- 
necting with  the  blepharoplast,  the  exact  limits  of  which  are  very  difficult 


Wilson,  M.     Spermatogenesis  in  the   Bryophyta.     Ann.   Bot,  25,   1911. 


52  DUDLEY    MEMORIAL   VOLUME 

to  determine.  The  cytoplasm  surrounding  the  nucleus  becomes  less  and 
less  evident  as  the  nucleus  increases  in  size  and  it  can  no  longer  be  clearly 
recognized  in  the  later  stages  of  development,  although  there  probably  per- 
sists a  thin  envelope  of  cytoplasm  surrounding  the  posterior  coils  of  the 
spermatozoid. 

The  blepharoplast  at  this  time  forms  a  short  hooped  prominence  at 
the  forward  part  of  the  spermatozoid  and  merging  insensibly  into  the  delicate 
cytoplasmic  prominence  which  extends  beyond  the  nucleus.  The  latter  con- 
tinues to  elongate  and  become  curved  over  in  the  plane  of  the  division  wall,  so 
that  the  older  spermatozoid  has  the  form  of  a  flat  coil.  (Fig.  7,  M.) 

When  fully  developed  the  sperm  is  a  slender  thread  composed  of  two 
complete  coils  and  part  of  a  third.  In  these  later  stages  the  double  stain 
of  safranin  and  gentian  violet  failed  to  clearly  differentiate  the  different  parts, 
the  spermatozoid  appearing  almost  uniformly  stained.  The  two  spermato- 
zoids  of  a  pair  are  very  closely  approximated  (Fig.  7,  M)  and  present  a 
very  characteristic  appearance. 

The  cilia  could  be  made  out  in  a  number  of  the  older  stages  (Fig.  7,  H), 
but  their  exact  origin  and  position  could  not  be  determined  as  accurately  as 
might  have  been  wished.  It  is  probable,  however,  that  as  in  some  other  cases 
which  have  been  investigated,  they  begin  to  double  up  at  an  early  stage  and 
arise  somewhat  back  of  the  apex.  Woodburn7  in  a  recent  paper  states  that 
in  Porella  the  anterior  end  of  the  spermatozoid  shows  a  slight  enlargement, 
which  he  interprets  as  the  blepharoplast.  The  cilia  in  this  case  arise  a  short 
distance  back  of  this  enlarged  part  of  the  blepharoplast.  No  trace  of  it  was 
observed  in  Calycularia. 

ARCHEGONIUM. 

The  female  plants,  as  we  have  already  seen,  are  decidedly  larger  than 
the  males,  and  are  usually  20-30  mm.  in  length,  with  a  breadth  of  from 
10-12  mm.  Like  the  male  plants,  they  are  usually  unbranched,  but  it  is 
not  uncommon  to  find  them  forked  once.  The  position  of  the  archegonia 
is  much  like  that  of  the  antheridia,  these  being  grouped  on  a  sort  of  recep- 
tacle. (Fig.  2,  A.)  As  a  rule,  only  one  archegonial  receptacle  is  formed, 
but  sometimes  the  thallus  will  resume  its  growth  and  a  second  one  may  be 
formed  near  the  apex.  The  archegonium  appears  to  agree  in  all  respects 
with  that  of  other  Jungermanniales  that  have  been  investigated.  (Figs.  8 
and  9.)  After  a  short  stalk  has  been  formed,  the  archegonium  mother  cell 
divides  by  the  usual  three  intersecting  vertical  walls  into  an  axial  cell  and 


Woodburn,  W.  L.     Spermatogenesis  in  Certain  Hepaticae.     Ann.  Bot.,  25,  1911. 


CALYCULARIA    RADICULOSA CAMPBELL 


53 


three  peripheral  ones.  From  the  axial  cell  is  cut  off  the  cover  cell  (Fig.  8, 
B  and  C)  and  then  follows  a  series  of  transverse  walls  separating  the  lower 
part  or  venter  from  the  upper  region  or  the  neck.  From  the  lower  of  the 
two  primary  axial  cells  the  egg  and  ventral  canal  cells  arise,  and  from  the 


Fig.  8.  A,  apex  of  female  plant,  showing  the 
position  of  the  archegonia,  x  about  40.  B-E, 
young  archegonia  in  median  section,  x  about  225. 

upper  ones  the  series  of  neck  canal  cells  and  the  outer  cells  of  the  neck. 
The  number  of  neck  canal  cells  is  variable,  but  to  judge  from  the  few  that 
were  examined,  there  are  first  formed  four  of  these  neck  cells,  some  or  all  of 
which  divide  again,  so  that  there  may  be  as  many  as  eight.  The  division, 
however,  is  very  often  not  complete  but  confined  to  the  nucleus  (Fig.  9,  C 
and  D ) .  As  usual  in  the  Jungermanniales,  the  neck  of  the  archegonium  has 
but  five  peripheral  rows  of  cells  (Fig.  9,  F). 


54 


DUDLEY    MEMORIAL    VOLUME 


In  the  ventral  region  peripheral  walls  occur  in  the  outer  cells,  so  that 
at  maturity  the  venter  is  more  or  less  completely  two  layered  ( Fig.  9,  D  and 
E). 

Figure  9,  G,  shows  an  abnormal  archegonium  from  a  receptacle  in  which 
one  of  the  archegonia  had  been  fertilized.  In  this  archegonium  there  were 
four  axial  cells,  all  of  which  were  a  good  deal  alike  and  all  except  one  much 
enlarged,  so  that  they  resembled  the  egg  of  the  normal  archegonium  more 
than  they  did  the  neck  canal  cells. 

/ 


Fig.  9.  A,  B,  two  sections  of  a  young  archegonium  with  five  neck  canal  cells; 
the  ventral  canal  cell  is  not  yet  formed,  x  about  225.  C,  a  somewhat  older  stage, 
showing  the  egg,  o,  and  the  ventral  canal  cell,  v.  D,  lower  part  of  a  nearly  ripe 
archegonium.  E,  full  grown  archegonium  which  has  failed  to  be  fertilized,  x  about 
90.  F,  cross-section  of  the  neck  of  archegonium.  G,  an  abnormal  archegonium 
with  unusually  large  axial  cells. 

After  fertilization  there  arises  about  the  group  of  archegonia  a  tubular 
envelope,  the  perianth,  which  finally  forms  a  very  conspicuous  vase-shaped 
sheath  around  the  sporogonium,  inside  the  involucre,  which,  like  that  of  the 
antheridial  receptacle,  is  made  up  of  very  much  laciniated  scales.  (Fig. 
1,  C.  D,  per.) 

THE  SPOROPHYTE. 

Only  a  few  very  young  embryos  were  found  and  these  were  not  well 
fixed,  so  that  it  was  impossible  to  make  a  satisfactory  study  of  the  develop- 
ment of  the  embryo.  The  youngest  sporogonia  of  which  successful  prepara- 


CALYCULARIA    RADICULOSA CAMPBELL 


55 


tions  were  made  were  already  far  advanced  and  were  differentiated  into 
the  various  parts. 

Even  before  the  first  division  takes  place  in  the  young  embryo,  the 
venter  of  the  archegonium  becomes  much  enlarged,  and  a  calyptra  is  devel- 
oped, enclosing  the  sporogonium  until  it  is  far  advanced.  This  attains  a 
thickness  of  5  or  6  layers  of  cells  at  the  base,  but  is  much  thinner  toward 
the  apex.  As  the  sporogonium  develops,  the  upper  portion  or  capsule 
becomes  oval  in  form,  and  below  consists  of  a  thick  seta,  which  terminates 
in  a  large  heart-shaped  foot  very  much  like  that  which  Cavers  describes  for 
Morkia  Flowtowiana.  (Cavers  Loc.  cit.  Fig.  37).  A  similar  foot  has  been 
observed  in  various  other  Jungermanniales. 


:B 


el 


Fig.  10.  A,  median  section  of  a  sporogonium  at  the  time  of  the  mitosis  of  the 
spore  mother  cells,  x  15.  B,  apical  region  of  a  younger  sporogonium,  sp.  spore 
mother  cells,  el.  young  elator,  x  225.  C,  part  of  the  lateral  wall  of  the  sporogonium. 

The  capsule  (Fig.  10)  has  a  relatively  thick  wall  which  is  better  de- 
veloped at  the  apex  than  at  the  sides,  this  difference  becoming  still  more 
marked  in  the  later  stages.  The  inner  tissue  now  shows  a  separation  into 
the  roundish  spore  mother  cells  and  the  elongated  young  elaters.  Long 
before  the  division  of  the  spore  mother  cells  begins,  they  show  the  first  indi- 
cations of  the  lobing  which  later  becomes  so  conspicuous.  At  this  stage  the 
walls  of  both  the  spore  mother  cells  and  the  elaters  are  very  delicate,  but 
can  be  readily  demonstrated  by  suitable  stains,  e.  g.  Bismarck  brown. 


56 


DUDLEY    MEMORIAL   VOLUME 


The  seta  at  this  stage  has  about  the  same  length  as  the  capsule,  and  in 
longitudinal  section  (Fig.  12,  H)  shows  the  cells  to  be  arranged  in  pretty 
regular  rows.  Probably  the  great  elongation  of  the  seta  at  the  time  the 
spores  are  shed  is  due  to  simple  elongation  of  the  cells  without  any  cell  divi- 
sions, as  has  been  shown  to  be  the  case  in  other  liverworts.  The  large  heart- 
shaped  foot  (f)  is  composed  of  somewhat  irregular  cells  showing  no  defi- 
nite arrangement. 


Fig.  11.  Spore  division.  A,  spore  mother  cell,  showing  the  quadripolar  spindle, 
x  750.  B,  three  spore  mother  cells  of  about  the  same  age,  showing  the  different 
arrangement  of  the  lobes  of  the  cell,  x  about  350.  C,  young  elater.  D,  spore  mother 
cell,  showing  the  chromosomes.  E,  a  somewhat  earlier  stage.  F-I,  successive  stages 
of  mitosis  with  quadripolar  spindle.  In  G  and  H  the  chromosomes  only  are  shown. 
/,  mother  cell  with  the  lobes  in  pairs;  there  are  two  nuclear  spindles  at  right 
angles  to  each  other.  K,  mother  cell,  showing  a  bi-polar  spindle  at  the  first  mitosis. 
L,  first  mitosis,  showing  two  nuclei  separated  by  a  distinct  cell-plate,  x  about  400. 
M,  mother  cell  just  before  the  final  separation  of  the  spores,  x  400. 


CALYCULARIA   RADICULOSA CAMPBELL  57 

THE  SPORE  DIVISION. 

(Figure  11.) 

One  sporogonium  showed  the  spore  mother  cells  in  process  of  division, 
all  stages  occurring  in  the  same  sporogonium.  The  preparation  was  some- 
what overstained  with  hsematoxylin,  but  nevertheless  showed  pretty  well  the 
details  of  division,  which  exhibited  a  considerable  amount  of  variation. 

The  spore  mother  cells  before  the  first  nuclear  division  are  deeply  four- 
lobed,  the  lobes  being  usually  arranged  tetrahedrally,  but  occasionally  placed 
in  pairs  at  right  angles  to  each  other  (Fig.  11,  J).  The  nucleus  is  large, 
but  in  this  over-stained  material  the  structure  was  not  usually  very  clear, 
the  nucleus  appearing  almost  homogenous.  A  few  specimens,  however, 
(Fig.  11,  E)  showed  a  more  or  less  granular  structure,  but  the  reticulum  was 
not  clearly  evident  nor  could  the  nucleolus  be  seen. 

A  striking  feature  was  noticed  in  most  of  the  cells,  viz. :  the  extension 
from  the  nucleus  into  each  of  the  four  lobes  of  the  cell  of  a  body  which  was 
apparently  the  same  as  the  "quadripolar"  spindle  described  by  Farmer  for 
Pallavicinia  decipiens*.  Often  the  center  of  each  lobe  was  occupied  by  a 
roundish  body  which  perhaps  marked  the  position  of  a  centrosome,  but  it 
cannot  be  stated  positively  that  centrosomes  are  present.  However,  as  cen- 
trosomes  occur  in  Pellia,  which  in  some  other  respects  suggest  a  relationship 
with  Calycularia,  it  is  possible  that  centrosomes  may  have  been  present  in 
this  case  also. 

In  the  later  stages  of  nuclear  division  some  differences  were  found  to 
occur.  Usually  the  process  seems  to  agree  very  closely  with  that  described 
by  Farmer  for  Pallavicinia  decipiens.  The  nuclear  membrane  disappears 
and  the  separate  chromosomes,  thick  oval  bodies,  can  be  made  out.  -(Fig. 
11,  F.)  There  are  eight  of  these  in  Calycularia  radiculosa  instead  of  the 
four  found  in  Pallavicinia  decipiens;  but  it  was  found  that  in  Pallavicinia 
radiculosa  there  were  also  eight,  as  there  are  in  Pellia. 

The  eight  chromosomes  next  divide  longitudinally  (Fig  11,  G),  and 
the  resulting  sixteen  chromosomes  separate  into  two  groups  (H).  Usually 
the  chromosomes  do  not  arrange  themselves  into  a  new  reticulum,  but  imme- 
diately undergo  a  second  division,  so  that  there  are  two  groups  of  16  chromo- 
somes (Fig.  11,  I),  each  of  which  separates  into  two  secondary  groups  of 
eight  chromosomes  which  finally  assume  the  form  of  resting  nuclei.  One 


8  Farmer,  J.   B.     Studies  in  Hepaticse  Pallavicinia  decipiens  Mitten.     Ann.  of 
Botany,  8,  35-52,  1894. 


58  DUDLEY    MEMORIAL    VOLUME 

of  these  nuclei  moves  to  each  lobe  of  the  mother  cell,  but  generally  remains 
near  the  center  of  the  .cell,  so  that  the  four  resulting  nuclei  are  quite  close 
together  (Fig.  11,  M). 

Not  all  of  the  spore  mother  cells,  however,  behave  in  this  fashion,  but 
sometimes  after  the  first  division  of  the  chromosomes  a  conspicuous  bi- 
polar spindle  of  the  usual  form  was  observed.  (Fig.  11,  K).  Later  two 
resting  nuclei  were  seen  with  a  cell-plate  between  them.  (Fig.  11,  L).  These 
secondary  nuclei  then  divided  again,  each  developing  another  bi-polar 
spindle,  these  secondary  spindles  sometimes  lying  at  right  angles  to  each 
other.  (Fig.  11,  J). 

After  the  nuclei  have  assumed  the  resting  condition  cell  walls  are 
formed,  simultaneosuly  extending  inward  from  the  indentations  between  the 
lobes  and  completely  dividing  the  mother  cell  into  its  four  component  parts, 
the  young  spores. 

The  ripe  spores  (Fig.  12,  F,  G)  possess  a  thick  membrane,  which  in 
sections  shows  two  well-marked  parts,  an  inner  uniform  layer  and  a  thick 
outer  one  provided  with  rounded  knobs,  which  give  it  a  very  characteristic 
appearance.  The  color  of  the  spore  is  dark  purple-brown,  like  the  thicken- 
ings upon  the  cells  of  the  capsule  wall.  It  is  probable  that  immediately 
adjacent  to  the  spore  cavity  is  a  thin  membrane  (intine)  of  cellulose,  but 
this  was  not  specially  investigated  and  did  not  show  clearly  in  the  sections 
that  were  examined.  The  nucleus  of  the  spore  is  rather  small  but  fairly 
conspicuous. 

The  elaters  (Fig.  12,  C,  D  and  E)  show  a  good  deal  of  variation. 
They  are  sometimes  very  much  attenuated,  with  the  spiral  bands  almost 
obliterated  at  the  ends,  suggesting  the  elaters  of  Makinoa,  where  the  spirals 
are  present  only  in  the  mid-region  of  the  elaters.  More  commonly,  however, 
they  taper  more  gradually  and  the  double  spiral  extends  to  the  end.  Con- 
siderable difference  in  size  may  be  noted  (Fig.  12,  E).  While  no  basal 
elaterophore  is  present,  occasionally  some  of  the  elaters  at  the  base  of  the 
capsule  seem  to  be  attached  at  one  end  and  suggest  a  rudiment  of  such  an 
elaterophore,  as  is  said  to  occur  in  the  other  species  of  Calycularia. 

The  surface  markings  of  the  spore  in  Calycularia  radiculosa  are  strik- 
ingly different  from  those  of  Pallavicinia  whether  of  the  section  Morkia  or 
Blyttia.  In  Pallavicinia  (See  Fig.  12,  K,  L)  the  surface  markings  have 
the  form  of  a  network  of  delicate  ridges,  such  as  also  occur  in  Fossombronia. 
This  marked  difference  in  the  character  of  the  spores,  together  with  certain 
other  differences,  might  be  considered  to  be  an  objection  to  uniting  Caly- 
cularia radiculosa  with  the  genus  Morkia. 

The  structure  of  the  capsule  wall  of  Calycularia  radiculosa,  according 


CALYCULARIA    RADICULOSA CAMPBELL 


59 


Fig.  12.  A,  apical  region  of  the  sporogonium  of  Calycularia  radiculosa,  showing 
the  thickenings  on  the  cell  walls,  x  90.  B,  lateral  wall  of  the  same  sporogonium. 
C,  ripe  spores  and  elaters,  x  about  200.  D,  very  much  attenuated  elaters,  x  350. 
E,  typical  elaters,  x  350.  F,  ripe  spore,  x  350.  G,  section  of  spore.  H,  lower  part 
of  seta  and  foot,  f,  x  90.  /,  apex  of  sporogonium  of  Pallavicinia  (Blyttia)  radi- 
culosa, x  90.  /,  lateral  wall  of  the  same,  x  90.  K,  section  of  ripe  spore,  x  350. 
L,  markings  on  the  surface  of  spore,  x  750. 


60  DUDLEY    MEMORIAL   VOLUME 

to  Schiffner,  is  Very  different  from  that  of  the  other  species  of  Calycularia. 
He  examined  C.  crispula  and  found  that  the  capsule  is  much  smaller  than 
in  C.  radiculosa,  and  was  perfectly  round  instead  of  being  oval.  The  wall 
showed  quite  a  different  structure,  being  composed  of  two  layers  of  cells  with 
somewhat  different  markings  from  those  found  in  Calycularia  radiculosa. 

In  the  ripe  sporogonium  in  the  latter  species  (See  Fig.  12,  A,  B)  the 
wall  is  much  thicker  at  the  apex,  where  there  are  five  or  six  layers  of  cells 
which  form  a  sort  of  apical  cap,  while  at  the  sides  there  are  usually  about 
four  layers  of  which  the  outer  one  is  composed  of  much  larger  cells,  the 
inner  layers  being  made  up  of  much  compressed  thin  walled  cells.  The 
radial  walls  of  the  outermost  layer  of  cells  are  marked  by  conspicuous  thick- 
ened bands,  which  are  sometimes  more  or  less  confluent,  giving  the  cells 
much  the  appearance  of  reticulate  tracheary  tissue.  The  inner  cells  have 
slight  thickenings,  but  these  are  very  irregularly  disposed,  and  are  almost 
wanting  upon  the  inner  cells  of  the  lateral  walls  of  the  sporogonium. 

At  maturity  there  may  be  recognized  four  valves  of  equal  size,  but  usu- 
ally these  do  not  separate  completely,  but  remain  together  in  pairs,  the  cap- 
sule opening  by  two  slits.  (Fig.  1,  D).  There  may  be  seen  between 
these  two  slits,  however,  a  delicate  line  marking  the  junction  between  the  two 
coherent  valves.  Schiffner  states  that  the  valves  never  separate  at  the  apex 
but  are  held  together  by  the  apical  cap  of  cells.  While  this  is  no  doubt  often 
the  case  (See  Fig.  2,  D),  it  may  happen  that  the  two  pairs  of  valves  separate 
completely.  (Fig.  E).  In  its  dehiscence,  therefore,  Calycularia  radiculosa 
is  more  like  Blyttia  than  it  is  like  Morkia.  In  the  other  species  of  Calycularia 
Schiffner  states  that  the  capsule  at  maturity  breaks  into  several  (5-6)  irregular 
parts  which  may  break  up  still  further,  thus  resembling  Fossombronia. 

The  seta  finally  becomes  very  long  ( Fig.  2,  D )  and  its  base  is  surrounded 
by  the  very  conspicuous  vase-shaped  perianth,  whose  opening  is  deeply  lobed 
and  fringed.  Material  of  Morkia  was  not  available  for  comparison  of  the 
structures  of  the  sporogonium  with  that  of  Calycularia  radiculosa,  but  sec- 
tions of  the  sporogonium  of  Pallavacinia  (Blyttia)  radiculosa  were  made. 
In  this  species  the  capsule  is  extremely  long,  cylindrical  and  very  little 
thicker  than  the  seta.  The  foot  is  pointed  and  not  clearly  delimited  from 
the  seta. 

The  apex  of  the  capsule  is  pointed  and  much  more  conspicuous  than  in 
Calycularia  (Fig.  12,  I).  The  capsule  wall  also  differs  in  the  character  of 
the  cells.  There  are  about  three  layers  of  cells  instead  of  four  and  the 
outer  cells  have  the  walls  uniformly  thickened  instead  of  showing  the  thick- 
ened bands  so  conspicuous  in  Calycularia  radiculosa.  (Fig.  12,  J.) 


CALYCULARIA    RADICULOSA CAMPBELL  61 

The  spores  are  very  different  in  their  markings,  as  we  have  already 
noted,  and  this  seems  to  be  .true  also  for  Morkia. 

THE  AFFINITIES  OF  CALYCULARIA  RADICULOSA. 

Schiffner,  from  his  study  of  Calycularia  radiculosa  concluded  that  it 
should  be  removed  from  its  present  association  with  C.  crispula,  C.  Birmensis 
and  C.  laxa  and  united  with  Morkia.  While  there  seems  to  be  reason  to 
remove  the  species  from  the  genus  Calycularia,  it  may  be  questioned  whether 
its  association  with  Morkia  is  justified.  While  Schiffner  states  that  Morkia 
is  without  conducting  tissue  in  the  mid-rib,  Cavers  has  shown  that  in  M .  Flo- 
towiana  there  are  two  strands  of  conducting  tissue,  but  in  Calycularia  radicu- 
losa these  are  entirely  wanting.  Moreover,  the  structure  of  the  sporogonium, 
i.  e.,  the  character  of  the  thickenings  of  the  wall  and  the  markings  of  the 
spores  are  quite  different,  and  more  like  Makinoa,  or  some  of  the  forms 
usually  referred  to  the  Codoniacese.  It  would  probably  be  better  to  con- 
sider this  plant  as  the  type  of  a  new  genus  intermediate  in  some  respects 
between  Morkia  and  some  of  the  less  specialized  forms  like  Makinoa  or 
Pellia.  It  is  hardly  likely  that  the  line  between  the  Codoniaceae  and  Lep- 
tothceas  (or  Blyttiaceae,  as  Cavers  has  called  them)  is  very  well  defined, 
and  it  is  probable  that  further  study  of  the  thallose  Jungermanniales  will 
result  in  decided  changes  in  the  accepted  arrangement  of  the  genera. 


STUDIES  OF  IRRITABILITY  IN  PLANTS. 
By  GEORGE  JAMES  PEIRCE,  Professor  of  Botany  and  Plant  Physiology. 

III. 

THE    FORMATIVE    INFLUENCE    OF    LIGHT. 

Introduction. 

IN  19061  I  published  a  paper,  under  the  above  title,  recording  the  results 
of  a  series  of  experiments  on  the  influence  of  the  direction  of  illumina- 
tion upon  the  shape  of  certain  plants.  The  most  striking  result  reported 
was  that  Anthoceros  plants  grown  from  the  spore  on  a  disc  revolving  in  a 
horizontal  plane,  and  therefore  receiving  fairly  equal  amounts  of  light  on 
all  sides  successively,  showed  no  trace  of  the  usual  dorsi-ventral  form  and 
structure  of  the  thallus  but  were  radial  in  structure,  cylindrical  or  conical 
in  form.  Anthoceros  jusijormis,  Aust.  and  A.  Pearsoni,  M.  A.  Howe,  both 
native  here  and  growing  within  a  short  distance  of  this  laboratory,  gave  the 
same  results;  but  the  spores  of  the  Marchantiaceous  liverwort  Fimbriaria 
(Asterella)  Californica  and  of  the  fern  Gymnogramme  triangularis  did  not, 
under  the  same  conditions,  give  rise  to  plants  round  in  section.  To  this 
extent  their  dorsi-ventrality  failed  to  show  itself — rhizoids  grew  equally  in 
all  directions  from  their  thalli  or  prothalli,  respectively — but  the  plants  were 
thin  plates,  though  curiously  crumpled,  as  the  figures  showed.  I  did  not 
understand  this  difference  in  result  and  have  tried  in  various  ways  to  ascer- 
tain the  reason  for  it.  I  have  not  yet  reached  a  satisfying  explanation,  but 
some  of  the  results  of  these  succeeding  experiments  are  interesting  enough 
to  record  now. 

THE  APPARATUS:  A  MULTIPLE  CLINOSTAT. 

The  apparatus  used  in  the  experiments  of  Czapek2,  which  suggested 
mine,  consisted  essentially  of  the  expensive  form  of  clinostat,  the  only  one 
generally  known  and  used  in  botanical  laboratories.  My  experiments  were 
carried  on  with  cheap  clocks,  modified  as  described  by  Ganong3.  Such  ap- 
paratus is,  however,  unreliable.  Indeed,  cheap  apparatus  may  be  the  most 
expensive.  Although  cheap  apparatus  may  perhaps  be  well  enough  for  an 


1  Annals  of  Botany,  XX,  449-465,  1906. 

2  Czapek,  F.  Weitere  Beitrage  zur  Kenntniss  der  geotropischen  Reizbewegungen. 
Jahrb.  f.  w.  Bot,  XXII,  261,  1898. 

3  Ganong,   W.   F.     A  laboratory  course  in   Plant   Physiology,   pp.    120-1,   New 
York,   1901. 


STUDIES    OF    IRRITABILITY    OF    PLANTS PEIRCE  63 

experiment  lasting  only  a  few  minutes  or  at  the  most  an  hour  or  two,  it 
ought  not  to  be  trusted  longer.  An  experiment  which  lasts  a  week  or  even 
months  increases  in  value  as  it  lasts,  if  it  had  any  value  at  the  start ;  and 
the  failure  of  the  apparatus  at  the  end  of  six  months  entails  a  loss  much 
greater  than  more  serious  mechanical  difficulty  within  a  day  or  two  of  the 
beginning.  The  cost,  unreliability  and  the  wearisomeness  of  winding  a  suit- 
able number  of  separate  instruments  drove  me  to  consider  other  apparatus. 
And  by  devising  new  apparatus  I  tried  to  ascertain  the  dominating  reason 
for  the  persistent  dorsi-ventrality  of  the  thalli  of  Fimbriaria  and  of  the  pro- 
thalli  of  Gymno gramme. 

To  Professor  W.  F.  Durand,  head  of  the  Department  of  Mechanical 
Engineering,  is  due  all  the  credit  for  structural  details  and  for  supervising 
the  construction  and  the  successive  modifications  of  the  apparatus  in  the 
Mechanician's  Shop  of  this  University.  And  I  take  this  opportunity  to  ex- 
press my  most  grateful  appreciation  of  his  skill  in  divining  what  I  wanted 
and  his  untiring  help  and  unflagging  patience  in  securing  it.  As  a  detailed 
description  of  the  apparatus  would  be  more  appropriate  elsewhere,  I  may 
here  content  myself  with  a  statement  of  its  essential  features. 

The  apparatus  may  be  called  a  multiple  clinostat.  As  my  experiments 
involved  the  turning  of  cultures  in  a  horizontal  plane  upon  a  vertical  axis, 
the  apparatus  began  with  a  set  of  twenty-five  turn-tables  on  five  shelves  built 
into  the  embrasures  of  each  of  three  windows,  the  turn-tables  in  each  window 
revolving  at  different  speeds,  but  all  the  turn-tables  in  one  window  turning 
at  the  same  rate.  The  actuating  mechanism  consisted  of  a  clock-work  driven 
by  a  heavy  weight  and  controlled  by  a  fan  governor.  This  actuating  mech- 
anism was  connected  by  a  series  of  belts  and  shafts  with  the  batteries  of  turn- 
tables. Experience,  however,  has  led  to  the  gradual  and  final  elimination 
of  all  belts.  Chains  and  sprockets  were  first  substituted  for  belts.  Finally 
these  were  replaced  by  direct  gears.  This  made  possible  the  consolidation 
of  the  cultures  into  one  window,  there  being  five  rows  of  ten  turn-tables  each 
on  a  set  of  shelves  in  the  window  nearest  the  clock-work  and  connected  with 
it  by  a  shaft  with  bevel  gears.  This  shaft  is  horizontal  and  runs  from  the 
clock-work,  bolted  to  a  table,  which  is  itself  bolted  to  the  floor,  to  a  vertical 
shaft  at  one  side  of  the  window.  At  each  shelf  this  vertical  shaft  carries  a 
gear  which  engages  a  corresponding  gear  carried  on  the  axle  of  the  nearest 
turn-table.  The  margin  of  this  turn-table,  and  of  all  the  others  in  the  row, 
is  toothed,  and  the  turn-tables  are  so  set  that  the  movement  of  one  of  them 
sets  all  the  others  into  similar  motion.  All  the  turn-tables  in  a  row  move  at 
the  same  rate,  but  the  rate  of  each  row  is  determined  by  the  ratio  of  the  gear 
on  the  vertical  shaft  to  that  of  the  first  turn-table  in  the  row.  It  is  possible, 


64  DUDLEY    MEMORIAL    VOLUME 

therefore,  to,  revolve  fifty  or  more  cultures  simultaneously,  but  at  five  differ- 
ent rates.  The  apparatus  runs  continuously,  day  and  night,  and  requires 
to  be  wound  only  once  in  thirty  hours. 

After  experience  and  reflection,  I  concluded  that  the  only  constant  and 
uniform  force  at  my  disposal  for  driving  my  clock-work  was  gravity.  This 
is  represented  by  discs  of  cast  iron  and  of  lead,  amounting  to  a  weight  of 
two  hundred  and  fifty  (250)  pounds — approximately  113.4  Kilos — which  is 
hung  from  a  pulley  running  on  a  wire  cable,  the  end  of  which  is  fastened  to 
a  beam  in  the  ceiling  of  the  room.  The  clock  is  wound  by  pulling  up  the 
weight  nearly  to  the  ceiling,  the  wire  winding  upon  a  drum  revolved  by  a 
crank. 

The  material  and  methods  of  culture  have  undergone  nearly  as  many 
changes  since  I  began  as  the  method  of  revolving  the  cultures.  With  the 
exception  to  be  noted  below,  I  still  use  small  crystallizing  dishes.  These  are 
about  7  cm.  diameter  and  3  cm.  depth.  I  have  had  small  square  tiles  of 
porous  flower-pot  clay  especially  made.  These  fit  into  the  bottom  of  the 
dishes,  and,  as  their  upper  surface  is  smooth,  the  least  possible  shadow  is  cast 
by  one  part  upon  another.  These  porous  tile  are  first  boiled  in  distilled 
water,  to  extract,  as  completely  as  may  be,  the  soluble  matter  which  they  may 
contain.  To  this  end  I  boil  the  tile  in  distilled  water  for  hours,  using  three 
or  four  waters  for  this  washing  and  leaching  process.  The  tiles  are  then 
allowed  to  drain  and  dry.  Meanwhile  Knopp's  Solution  is  made  as  follows : 

SOLUTION  A  SOLUTION  B 

KNOs    2grs  Ca(NO3)2   8  grs 

MgSO4     2  "  Aq.  dist 3000  cc 

KsHPCU    2  « 

Aq.  dist 1000  cc 

To  1  part  of  A  3  parts  of  B  are  added  and  the  mixture  boiled  for  fifteen 
minutes  in  a  cotton-plugged  flask.  The  resulting  precipitate  is  either  filtered 
off  or  allowed  to  settle  in  the  bottom  of  the  flask.  At  all  events,  the  clear 
liquid  is  poured  into  the  culture  dishes  to  about  three-quarters  the  depth  of 
the  porous  tiles.  The  dishes  are  covered  with  the  lids  or  bottoms  of  Petri 
dishes  of  suitable  size.  They  are  now  steam  sterilized  for  an  hour  or  more 
and  are  allowed  to  cool  over  night  in  the  sterilizer.  As  it  is  necessary  to 
prevent  light  from  falling  otherwise  than  from  the  side  upon  the  plants  to  be 
cultivated,  the  lids  of  the  dishes  are  given,  after  cooling,  a  smooth  continuous 
coat  of  dull  black  "Japalac,"  an  inexpensive  and  quickly  drying  varnish  easily 
applied. 

It  may  be  wondered  why  I  did  not  use  Plaster  of  Paris  poured  into  the 
crystallizing  dishes  and  allowed  to  set  in  them  as  molds.  These  would  have 


STUDIES    OF    IRRITABILITY    OF    PLANTS PEIRCE  65 

furnished  a  fairly  smooth  substratum  of  convenient  extent.  Two  considera- 
tions prevented,  namely  :  the  solubility  of  Plaster  of  Paris,  and  its  color.  The 
latter  could  have  been  modified  by  lamp-black  or  any  other  insoluble  pigment, 
but  the  constant  presence  of  an  undue  amount  of  calcium  sulphate  in  the 
water  which  the  plants  were  to  absorb  seemed  to  me  unnecessary  and  possibly 
confusing.  Flower-pot  tile  is  certainly  more  nearly  like  soil  in  color  and 
composition  than  is  Plaster  of  Paris,  more  convenient  to  handle,  and  readily 
enough  obtained  in  any  desired  size,  if  a  reasonable  number  of  tiles  be  ordered 
at  any  one  time. 

The  spores  are  sowed  as  uniformly  as  possible  on  the  now  damp  tile, 
which  is  standing  in  sterilized  Knopp's  Solution  in  the  dishes.  The  spores 
are  sowed  from  stiff  smooth  writing  paper  by  tapping  the  paper  with  a  pencil 
or  paper  knife  in  such  a  way  as  to  discharge  a  fairly  even  shower  of  spores 
upon  the  tiles.  The  culture  dishes  are  then  marked  and  put  in  place  on  the 
turn-tables,  and  on  the  shelf  beside  them  as  controls,  respectively. 

The  speed  of  the  turn-tables  is  a  matter  of  considerable  importance. 
The  greater  the  speed,  the  greater  the  amount  of  power  required.  In  no  case, 
however,  have  I  used  a  speed  at  which  the  centrifugal  force,  even  at  the  edge 
of  a  culture,  could  have  had  any  part  in  the  result ;  and  at  the  center  of  revo- 
lution, which  is  also  the  point  of  most  nearly  equal  illumination,  there  would 
be  no  centrifugal  force.  The  speeds  which  I  have  so  far  used,  in  addition 
to  those  previously  reported4,  are  the  following : 

10  turn-tables  making  four  revolutions  a  minute. 
20  "  "        two  "  "       " 

10  "  "        one 

I  have  also  arranged  to  have  a  fifth  row  of  ten  turn-tables,  so  geared  as 
to  make  a  complete  revolution  in  two  minutes.  I  may  add  that,  although 
I  have  so  far  used  the  turn-tables  only  on  vertical  axes,  I  have,  nevertheless, 
had  the  shelves  so  attached  to  the  frame  of  the  shelving  in  the  window  that 
they  may  be  set  at  any  desired  angle  between  the  vertical  and  the  horizontal. 
The  positions  and  structures  of  the  gears  of  the  vertical  distributing  shafts 
and  of  the  first  turn-tables  in  each  row  must  be  and  may  be  modified  accord- 
ingly. Obviously,  if  the  turn-tables  are  to  be  used  in  a  position  in  which 
their  axes  would  point  obliquely  downward,  it  would  be  necessary  to  use  cups, 
into  which  the  axes  could  be  locked ;  but  for  my  experiments  so  far,  no  modi- 
fication of  the  cups  has  been  necessary.  The  cups  carrying  the  axes  of  the 
turn-tables  are  cast  steel  with  a  steel  ball  of  suitable  size  in  the  bottom  of 


Annals  of  Botany,   XX,   1906. 


66  DUDLEY    MEMORIAL   VOLUME 

each  cup,  and  screwed  to  the  shelves.  So  long  as  the  diameter  of  the  bore 
of  the  cups  remains  uniform  and  the  wear  of  the  axis  corresponds,  the  motion 
of  the  turn-tables  should  be  uniform.  The  turn-tables  themselves  are  made 
of  a  light  alloy,  Alzine,  which  sometimes  appears  to  be  too  brittle;  but  if 
not,  teeth  of  great  uniformity  may  be  cut  in  the  edges  of  the  turn-tables. 
When  the  regular  teeth  of  adjacent  turn-tables  are  so  set  that  they  do  not 
bind  or  allow  too  much  play,  with  the  inevitable  contraction  and  expansion 
of  the  shelves  and  frame  in  the  changing  temperatures  and  humidities  of  a 
laboratory,  the  clock-work  drives  them  with  great  regularity.  Indeed,  next 
to  the  very  desirable  feature  of  carrying  many  cultures  at  once  on  this  multi- 
ple clinostat,  the  regularity  of  revolution  is  its  most  valuable  feature. 

It  may  not  be  necessary  to  add  that  the  multiple  clinostat  now  in  my 
laboratory,  and  thus  briefly  described,  is  the  product  of  the  experiments,  fail- 
ures and  successes,  of  the  last  six  years.  Each  improvement  has  been  the 
fruit  of  failure.  Some  of  these  failures  have  been  very  disheartening,  for 
one  does  not  like  to  lose  or  to  vitiate  the  accumulated  result  of  six  or  eight 
months  of  work  by  the  clock-work  or  any  set  of  turn-tables  coming  to  a  stand- 
still for  an  hour. 

EXPERIMENTAL    WORK. 

Only  one  or  two  of  the  questions  suggested  by  my  previous  work  and 
left  unanswered  in  that  paper  will  be  considered  in  this.  The  plants  which 
I  have  experimented  upon  have  been  the  prothalli  of  Pteris  aquilina  and  Gym- 
nogramme  triangularis,  grown  from  the  spore  on  tile;  plants  of  Porella  Bo- 
landeri,  a  foliose  liverwort  which  I  brought  into  the  laboratory  from  rocks 
and  tree-trunks  near  by  and  cultivated  on  the  tile  in  crystallizing  dishes; 
plants  of  Fimbriaria  (Asterella)  Caltfornica,  also  grown  from  the  spore ; 
Anthoceros  fusiformis,  grown  from  the  spore  and  used  simply  as  a  check,  for 
the  results  on  turn-table  and  shelf  were  the  same  as  previously  reported;  and 
plants  of  white  mustard  and  of  wheat,  raised  in  two-inch  flower  pots  in  good 
soil  from  the  seed.  The  results  are  in  the  main  similar,  and  I  shall  discuss 
them  all  together  after  separately  describing  the  experiments  on  the  different 
sorts  of  plants. 

Porella  Bolanderi  (Aust.)  Pearson. 

On  November  14,  1907,  I  collected  plants  of  Porella  Bolanderi  growing 
on  rocks  and  tree-trunks  about  a  half  mile  from  this  University.  The  plants 
were  dry  and  dormant.  I  sorted  these,  after  moistening  with  sterilized  water, 
and  selecting  clean  branches  about  a  centimeter  long,  placed  these  upon 


STUDIES    OF    IRRITABILITY    OF    PLANTS PEIRCE  67 

sterilized  tiles  in  crystallizing  dishes,  containing  sterilized  Knopp's  Solution 
of  0.35%  concentration  and  covered  with  blackened  lids.  These  dishes  I 
set  on  the  turn-tables  and  on  the  shelf  beside  the  turn-tables.  I  took  all 
possible  pains  to  select  branches  clean  and  healthy-looking,  but  as  no  steriliza- 
tion of  the  material  was  possible,  it  was  inevitable  that  a  certain  amount  of 
moulding  should  take  place.  All  of  the  cultures  succumbed  to  infections 
sooner  or  later,  but  enough  of  them  grew  well  to  justify  a  record  of  the  experi- 
ment, although  I  do  not  by  any  means  regard  it  as  concluded. 

The  general  characteristics  of  this  plant  are  well  known.  Detailed  de- 
scriptions may  be  found  in  Campbell's  "Mosses  and  Ferns"5  and  elsewhere. 
For  our  purpose  it  is  sufficient  to  say  that  the  plant  is  dorsi-ventral  to  the 
extent  of  having  two  sets  of  leaves :  foliage  leaves,  which  are  green  and  closely 
arranged,  forming  the  upper  side  of  the  plant,  and  the  so-called  amphigastria, 
leaves  or  scales  not  green,  and  overlapping  along  the  under  side  of  the  stem. 
The  plant  grows  more  or  less  closely  appressed  to  the  sub-stratum,  whether 
this  is  vertical,  oblique,  or  horizontal ;  that  is,  the  plant  grows  at  right  angles 
to  the  direction  from  which  most  of  the  light  comes.  If,  therefore,  the  plant 
be  put  on  a  horizontal  surface  and  the  light  be  made  to  fall  more  or  less 
horizontally  upon  it,  the  plant  or  its  leaves  should  so  turn  that  its  foliage 
leaves  would  stand  mainly  at  right  angles  to  the  incident  rays,  and  the  amphi- 
gastria should  be  on  the  side  away  from  the  light.  This  happens  with  the 
plants  growing  on  the  shelf,  receiving  light  always  in  one  and  the  same  direc- 
tion, just  as  it  would  happen  in  the  case  of  a  Porella  plant  growing  from  a 
horizontal  to  a  more  or  less  vertical  sub-stratum  out  of  doors.  The  case  of 
the  plants  on  the  turn-tables,  on  the  other  hand,  is  quite  different,  for  they 
have  no  darker  side.  The  position  of  all  the  leaves  and  of  the  amphigastria 
on  the  older  parts  of  these  plants  changes ;  they  flare  more  from  the  stem. 
On  the  younger,  as  well  as  on  the  older  parts,  the  amphigastria  become  less 
scale-like  and  grow  more  and  more  leaf-like.  I  have  no  doubt  that  this  ex- 
periment, continued  with  greater  freedom  from  infections  than  I  secured  in 
1907,  would  yield  results  entirely  similar  to  those  of  Nemec6,  but  I  have  not 
yet  been  able  to  repeat  it  and  carry  it  through.  I  would  suggest  here  only 
that  experiment  seems  likely  to  confirm  the  opinion  of  morphologists  that 
amphigastria  are  modified  leaves,  and  to  show  that  they  develop  as  they  do 
partly  because  they  are  on  the  shaded  parts  of  these  plants. 


5  Campbell,    D.    H.     The    Structure   and    Development   of   Mosses   and    Ferns. 
2d  Edition,  New  York,   1905. 

6  Nemec,  B.    Die  Induktion  der  Dorsiventralitat  bei  einigen  Moosen.     Bull.  int. 
de  1'Acad.  Sci.  de  Boheme,  1904,  1906. 


68  DUDLEY    MEMORIAL   VOLUME 

Fimbriaria  (Asterella)  Calijornica,  Hampe. 

Spores  of  this  plant  were  collected  at  the  beginning  of  the  dry  season, 
late  in  April  or  in  May,  according  to  the  time  of  ripening,  from  plants  grow- 
ing on  a  sandy-loam  bank,  not  far  from  the  laboratory.  The  spores  were 
kept  in  envelopes  and  pasteboard  boxes  in  a  case  in  the  laboratory  and  were, 
therefore,  air-dry  and  well  ventilated.  As  the  humidity  of  the  air  runs  low 
during  the  summer  dormant  period,  both  in  the  laboratory  and  out  of  doors, 
the  spores  were  necessarily  inactive  for  months.  With  greater  humidity  or 
— what  would  produce  this — inferior  ventilation,  their  respiratory  activities7 
would  be  greater  and  there  would  be  danger  that  their  germinating  and 
other  powers  might  be  impaired.  For  this  reason  I  avoid  keeping  spores  or 
seeds  which  are  to  be  used  for  germination  in  tightly-closed  jars  or  bottles. 
If  it  be  necessary  to  protect  them  against  mice,  I  use  tins,  the  lids  of  which 
close  them  loosely  enough  to  permit  more  or  less  circulation  of  air.  In  this 
way  spore  and  seed  deterioration  is  delayed  and  normal  dormancy  is  main- 
tained. 

The  spores  were  sowed,  as  above  described,  on  sterilized  tile  in  black- 
covered  crystallizing  dishes,  on  October  11,  1911.  Of  these,  five  were  put 
on  turn-tables  making  two  complete  revolutions  per  minute,  and  three  were 
set  on  the  shelf  beside  them.  The  crystallizing  dishes  standing  on  the  shelf 
were  marked  on  the  side  away  from  the  window,  so  that  one  might  always 
know  the  original  exposure  and  the  more  easily  maintain  it. 

As  previously  shown8,  the  direction  of  the  plane  of  division  in  the  ger- 
minating spores,  and  of  growth  in  the  germ-tube,  is  determined  by  the  direc- 
tion from  which  the  light  falls  upon  the  spores.  On  the  clocks,  therefore, 
the  spores  germinate  in  every  direction,  and  the  plantlets  are  erect  from  the 
start.  Germination  actually  begins  almost  at  once,  no  doubt,  but  the  evi- 
dences of  it  are  plainly  visible  within  ten  days  after  sowing  the  spores.  In 
the  shelf  cultures  the  plantlets  are  prostrate,  growing  toward  the  light  as 
single  chains  of  chlorophyll-containing  cells.  In  these  latter  cultures,  as  in 
nature,  the  light  falls  upon  the  plantlets  mainly  from  one  direction,  and  the 
plantlets  react  accordingly.  After  the  plantlet  has  become  a  single  chain 
of  several  cells,  the  end  cell  repeatedly  divides  in  such  planes  as  to  change 
the  plantlet  to  a  conical  shape.  These  little  cones,  with  their  apices  pointing 
away  from  the  light  and  obliquely  downward,  grow  both  in  length  and  in 
diameter  fairly  symmetrical  for  a  few  weeks.  After  a  time,  however,  they 


7  Babcock,  S.  B.     Metabolic  water ;  its  production  and  role  in  vital  phenomena. 
Research   Bull.  22,   Univ.   Wis.   Agric.   Exp.   Sta.,   March,   1912. 

8  Peirce,  G.  J.    Annals  of  Botany,  XX,  p.  453+,  1906. 


STUDIES    OF    IRRITABILITY    OF    PLANTS PEIRCE  69 

cease  to  be  symmetrical,  the  side  of  the  cone  away  from  the  light  growing  out 
in  a  form  more  or  less  shelf-like.  This  is  the  beginning  of  the  thallus  of  the 
more  or  less  mature  form. 

In  cultures  I  have  never  carried  the  plants  beyond  this  beginning — six 
months  or  thereabouts  after  sowing  the  spores — for  although  plants  of  the 
same  species  normally  survive  the  summer  out  of  doors9,  they  do  not  with- 
stand the  much  more  complete  drying  in  a  culture  dish,  or,  if  an  attempt  is 
made  to  keep  them  moist  over  summer,  they  succumb  to  fungus  enemies.  I 
do  not  know  what  would  happen  if  the  cultures  were  kept  continuously  on  the 
shelves  and  on  the  revolving  turn-tables,  for  it  has  never  been  possible  for  me 
to  stay  in  my  laboratory  throughout  the  long  summer  vacation,  and  I  have 
so  far  been  unable  to  arrange  to  have  my  clock-work  regularly  wound,  i.  e., 
daily  throughout  my  absences.  It  is  usually  easier,  in  a  laboratory  as  well 
as  elsewhere,  to  provide  apparatus  than  to  secure  assistants.  Hence,  at  the 
end  of  the  college  year,  in  May,  I  am  obliged  to  take  my  cultures  from  their 
places  on  shelf  and  turn-table  and  set  them  away  in  a  dark  cupboard.  They 
remain  there  till  September.  During  these  months  they  have  succumbed  to 
mould  or  drought.  Some  day,  however,  I  shall  be  able  to  carry  them  along 
continuously  from  the  spore  to  the  production  of  spores  again. 

Turning  now  once  more  to  the  plantlets  subjected,  on  the  turn-tables, 
to  light  from  all  directions  successively,  we  find  that  they  maintain  the  erect 
position  which  they  assume  immediately  on  germination.  They  thicken  at 
the  ends  away  from  the  spores,  and,  since  they  are  revolved  in  a  horizontal 
plane  and  receive  light  mainly  horizontally,  they  become  vertical.  I  do  not 
think  the  force  of  gravity,  or  the  presence  of  water  below  them,  or  any  othei 
influence  than  light  has  much  to  do  with  the  erect  position  of  these  plantlets. 
They  become  erect  cones  standing  on  their  apices,  and,  uniformly  on  all 
sides,  they  develop  rhizoids,  which  attach  them  to  the  tiles.  The  little  plants 
are  thus  stayed  and  kept  from  toppling  over.  They  keep  pace  in  their  growth 
with  the  plants  receiving  light  from  one  side  only,  on  the  shelf,  and  after  a 
time  exceed  them  considerably  in  size.  On  account  of  the  difficulties  pre- 
viously enumerated,  however,  I  have  never  been  able  to  carry  these  plants 
through  the  summer  or  continue  the  experiment  for  more  than  seven  months. 
Though  the  plants  on  the  shelf  are  at  first  and  for  some  weeks  radial  in 
structure,  they  sooner  or  later  go  over  to  the  dorsi-ventral  form  under  the 
influence  of  light  falling  upon  them  from  one  direction  only.  This  form 
they  maintain  throughout  all  but  the  very  early  stages  of  their  existence. 
The  plants  revolving  on  the  turn-tables,  on  the  other  hand,  remain  radial  in 


9  Campbell,  D.  H.     Resistance  of  drought  by  liverworts.     Torreya,  IV,  1904. 


70  DUDLEY    MEMORIAL    VOLUME 

structure,  conical  in  form,  for  a  much  longer  time,  and  in  so  doing  resemble 
Anthoceros™.  For  the  reasons  given  above  I  cannot  say  that  they  would 
always  maintain  this  radial  structure,  though  experience  with  these  plants 
and  with  Anthoceros  leads  me  to  believe  that  dorsi-ventrality,  in  these  two 
genera  at  least,  is  not  alone  inherited,  but  that  it  is  a  product  of  circumstances 
as  well  as  of  substance,  the  continuity  of  substance  and  the  continuity  of 
influence  (direction  of  illumination)  from  generation  to  generation  insuring 
the  repetition  of  this  quality  in  successive  generations. 

It  may  be  suggested  that  the  dorsi-ventrality  does  not  develop  in  the 
plants  revolving  on  the  turn-tables,  not  because  it  is  not  inherited,  but  because 
it  is  prevented  from  appearing  because  one  of  the  conditions  for  its  devel- 
oping is  lacking.  I  do  not  care  to  contribute  to  a  revival  of  the  profitless 
discussion,  wisely  dropped,  involving  a  conceivable  if  unnecessary  distinction 
between  condition  (Bedingung)  and  stimulus  (Reiz),  for,  as  will  more 
plainly  appear  in  the  next  section  (pp.  70-74),  the  influence  of  the  direction 
of  illumination  is  active  rather  than  passive.  When  one  compares,  at  the 
end  of  an  experiment  which  has  lasted  for  months,  the  sizes  of  the  plants  on 
the  turn-tables  with  those  on  the  shelf,  one  realizes  the  greater  size  of  those 
more  uniformly  illuminated,  symmetry  and  size  going  together. 

The  result  of  growing  Fimbriaria  Calif  arnica  from  the  spore  under  con- 
ditions of  equal  and  of  unequal  illumination  from  all  directions  successively, 
is  the  same,  so  far  as  the  experiments  could  be  continued,  as  with  the  two 
species  of  Anthoceros  previously  reported  upon,  but  the  longer  life-cycle  of 
Fimbriaria  makes  it  necessary  to  continue  the  experiment  for  a  longer  time 
than  has  so  far  been  possible,  in  order  to  reach  a  definitive  result  and  to 
justify  a  final  conclusion.  Fimbriaria  grown  from  the  spore  does  not  fruit, 
at  least  in  my  cultures,  within  the  time  limits  of  one  natural  growing  season — 
that  is,  between  the  first  rains,  say  early  in  November,  and  the  beginning  of 
the  dry  season,  in  May.  On  the  other  hand,  Anthoceros  does,  but  its  spores 
are  not  equally  fertile  in  successive  seasons,  and  since  the  wonderful  crop  of 
1905  I  have  been  unable  to  secure  spores  of  such  vigor  that  I  cared  to  con- 
tinue experimenting  upon  the  plantlets  beyond  confirming  previous  results. 
Nor  is  it  necessary,  as  the  results  reported  in  the  next  section  will  show. 

Pteris  aquilina  and  Gymnogramme  triangularis. 

Spores  of  Pteris  collected  in  southern  California  in  June,  and  of  Gym- 
nogramme collected  near  the  laboratory  in  September,  were  sowed  on  October 
11,  1911,  under  the  conditions  previously  described.  Six  of  the  cultures  of 
Pteris  were  set  on  turn-tables  revolving  four  times  a  minute,  two  on  the  shelf 

10Peirce.     Ann.  Bot.,  1906. 


STUDIES    OF    IRRITABILITY    OF    PLANTS PEIRCE  71 

by  them.  A  second  sowing  in  other  dishes  was  made  on  November  13,  1911. 
About  the  same  number  of  cultures  of  Gymnogramme  were  started  on  Octo- 
ber 11  and  November  13.  The  results  were  similar  to  those  obtained  with 
Pteris  but  less  striking,  because  of  the  smaller  size  of  the  prothalli.  Further- 
more, the  Pteris  material  was  so  much  freer  from  contaminating  blue-green 
algae  that  the  cultures  were  correspondingly  more  satisfactory.  Since  steriliz- 
ing the  spores  is  impossible,  it  is  not  usually  possible  to  make  a  pure  culture 
of  fern  prothalli  directly  from  the  spore. 

The  spores  of  both  ferns  germinated  well,  behaving  during  the  earlier 
stages  on  the  turn-tables  and  on  the  shelf  as  before  described11.  In  the  ex- 
periments previously  reported,  the  spores  of  Gymnogramme  germinating  on 
turn-tables  developed  into  thin  prothalli,  crumpled  or  waved  instead  of  flat, 
heart-shaped  instead  of  conical  or  cylindrical,  but  with  rhizoids,  antheridia 
'and  archegonia  in  equal  numbers  on  both  sides,  and  with  the  prothalli  stand- 
ing erect,  but  at  all  possible  angles  on  the  tiles.  In  the  shelf  cultures,  on  the 
other  hand,  the  prothalli  were  normal  in  shape  and  almost  linearly  placed,  in 
ranks  surprisingly  regular,  at  right  angles  to  the  incoming  light  from  the 
window.  These  results  I  thought  might  be  due  to  the  slow  revolution  of  my 
turn-tables  and  I  hoped  by  using  quicker  ones  I  might  obtain  cylindrical 
or  conical  prothalli.  I  have  not  yet.  Why,  I  do  not  know.  It  may  be  that 
the  turn-tables,  revolving  four  times  a  minute  instead  of  once  in  fifteen  as 
before,  are  still  too  slow,  or  it  may  be  that  continuous  as  well  as  uniform 
illumination  is  necessary. 

The  results  of  five  months'  growth  on  turn-tables  are  shown  in  Figures 
1  and  2.  Figures  3  and  4  show  shelf  cultures.  These  two  sets  of  photo- 
graphs of  Pteris  cultures  sowed  on  October  11,  1911,  were  kindly  taken  for 
me  by  Mr.  James  McMurphy,  Instructor  in  Botany  in  this  University.  The 
culture  dishes  were  uncovered  and  placed  on  the  horizontal  stand  of  a  verti- 
cally working  camera,  so  focused  as  to  give  a  picture  double  natural  size.* 
The  magnification  was  the  same  for  all  five  photographs.  This  was  made 
possible  by  the  uniform  thickness  of  the  tiles  and  of  the  glass  of  the  dishes 
The  four  figures  are,  therefore,  perfectly  comparable. 

Figure  1  shows,  within  the  circular  line  which  indicates  the  bottom 
of  the  crystallizing  dish,  and  upon  a  square  porous  tile,  the  corners  of  which 
have  been  knocked  off  to  fit  the  glass  dish,  a  large  number  of  fern  prothalli 
of  two  very  different  sizes.  These  prothalli  are  erect  or  nearly  so.  Those 
nearest  the  center  of  the  tile  (the  center  of  revolution)  are  most  nearly  erect 
and  most  plainly  show  the  copious  growth  of  rhizoids  on  both  sides.  The 

«  Annals  of  Botany,  XX,  p.  454+,  1906. 
*  Reduced  to  natural  size  in  the  figures. 


72  DUDLEY    MEMORIAL    VOLUME 

larger  prothalli  bear  archegonia,  the  smaller  prothalli  are  antheridial. 
Toward  the  upper  right  hand  corner  is  a  young  sporophyte,  showing  that 
conditions  in  the  culture  were  sufficiently  favorable  to  permit  fertilization 
and  subsequent  development.  Microscopic  examination  shows  that  the  repro- 
ductive organs  are  borne  on  both  sides  of  the  prothallus,  as  uniformly  as  the 
rhizoids.  The  prothallus  has  the  usual  cushion,  which  bears  the  rhizoids, 
archegonia  and  antheridia. 

Near  the  lower  edge  of  the  figure  is  a  large  archegonial  prothallus, 
irregular  in  outline  but  plainly  dorsi-ventral  in  structure.  This  has  rhizoids 
only  on  the  side  toward  the  middle  of  the  tile.  Comparing  this  prothallus 
with  the  two  nearest  the  center  of  the  culture,  one  sees  that  the  more  uniform 
illumination  of  the  plants  in  the  middle  is  accompanied  by  a  more  uniform 
growth  of  rhizoids.  The  distance  between  the  prothallus  at  the  edge  and 
those  in  the  middle  of  the  tile  is  about  2  centimeters,  but  this  slight  difference 
in  position  is  accompanied  by  enough  difference  in  illumination  to  permit  the 
plantlet  near  the  edge  to  complete  its  usual  dorsi-ventral  development,  while 
the  plants  at  the  center  form  rhizoids  and  reproductive  organs  equally  on 
both  sides. 

From  this  one  may  infer  either  that  the  fern  prothallus  is  very  sensitive 
to  light,  since  slight  differences  in  illumination  cause  such  evident  differences 
in  behavior,  or  that  it  is  only  slightly  sensitive,  since  exactly  uniform  illumina- 
tion is  necessary  to  overcome  the  usual  dorsi-ventrality  in  any  degree. 
Whether  the  plant  is  sensitive  or  not  sensitive  can  be  proved  only  by  experi- 
ment. One  of  these  experiments  would  involve  constant  illumination,  and 
this  I  hope  to  try  shortly. 

In  Figure  2  the  number  is  larger,  the  distribution  more  regular,  and  the 
results  striking. 

Figure  3  is  that  of  a  shelf  culture,  the  direction  of  illumination  of  which 
is  indicated  by  the  arrow.  The  size,  position,  form  and  appendages  of  the 
largest  archegonial  prothalli  are  quite  as  usual  and  normal.  Rhizoids  and 
archegonia  develop  only  on  the  side  away  from  the  light,  the  prothalli  are 
erect  in  response  to  the  nearly  horizontal  plane  along  which  the  light  is  re- 
ceived, but  they  are  nearly  flat  and  extend  at  right  angles  to  the  light  and 
they  are  of  the  usual  size.  The  much  (and  normally)  smaller  antheridial 
prothalli  correspond  in  all  these  respects  with  the  much  more  striking  arche- 
gonial. 

I  do  not  know  what  may  be  said  to  be  the  normal  ratio  between  arche- 
gonial and  antheridial  prothalli  in  Pteris,  and  I  think  most  botanists  would 
doubt  there  being  any  "normal"  apart  from  the  conditions  or  circumstances 
in  nature  or  in  an  experimental  culture,  but  the  ratio  of  archegonial  prothalli 
to  antheridial  in  the  two  turn-table  cultures  figured  is  certainly  larger  than 


STUDIES    OF    IRRITABILITY    OF    PLANTS PEIRCE  73 

in  the  shelf  culture  shown  in  Figure  3.  Figure  4  shows  another  shelf  culture, 
but  this  was  robbed  from  time  to  time  of  some  of  its  prothalli,  both  arche- 
gonial  and  antheridial,  for  purposes  of  examination.  Pains  were  taken,  how- 
ever, to  remove  such  numbers  of  male  and  female  prothalli  as  to  maintain 
the  ratio.  We  see  that  this  ratio  is  quite  different  in  the  shelf  and  in  the 
turn-table  cultures,  there  being  a  larger  proportion  of  female  prothalli  in  the 
turn-table  cultures. 

Measurements  of  typical  archegonial  prothalli  from  turn-table  and  shelf 
cultures  were  very  kindly  made  for  me  by  a  student  in  my  laboratory,  Miss 
Viola  F.  Nichols,  whom  I  wish  to  thank  for  her  help.  The  data  follow : 

PTERIS  sowed  X,  11,  '11.  SHELF  TURN-TABLE 

measured  IV,  15,  '12.       8x4  mm.  10  x  5  mm. 

6x4  11  x  5 
5  x  3.5  9x5 

5x3  10  x  4 

2.5  x  1  10  x  5 
4  x  2.5  8x4 

4x3  8  x  4.5 


average'  4.85  x  2.85  mm.  9.42  x  4.64  mm. 


measured  IV,  15,  '12 

5x3 

10  x  6 

10  x  4 

10  x  4 

9x4 

8x4 

7x4 

average 

5x3  mm. 

9.00  x  4.33  mm. 

mean  of  2  averages 

4.91  x  2.91 

9.21  x  4.48 

"      area 

14.28  sq.  mm. 

41  .  26  sq.  mm. 

ratio  of  areas 

1 

to             2.8 

The  difference  in  size,  so  apparent  to  the  eye,  is  thus  confirmed  by  meas- 
urements. Accompanying  this  difference  in  size  is  a  more  than  correspond- 
ing difference  in  the  number  of  archegonia  and  antheridia  on  the  turn-table 
prothalli  as  compared  with  those  in  the  shelf  cultures.  This  ratio  is  nearer 
four  or  five  to  one.  For  this  figure  I  am  also  indebted  to  Miss  Nichols,  but 
my  own  observations  correspond. 

These  figures,  together  with  an  inspection  of  the  photographs,  furnish 
the  evidence  of  differences  between  Pteris  prothalli  grown  on  turn-tables  in 
approximately  uniform  illumination  on  all  sides  successively,  and  others 
grown  on  the  shelf  with  the  light  always  from  one  side.  These  differences 
are  of  three  main  sorts:  1st,  in  size  of  the  vegetative  parts,  the  prothalli; 
2nd,  the  numbers  of  reproductive  organs;  3rd,  the  proportions  of  male  and 


74  DUDLEY    MEMORIAL   VOLUME 

female  prothalli.  These  axe  very  surprising,  but  what  I  have  recorded  above 
for  Pteris  aquilina  sowed  X,  11,  1911,  is  equally  true  of  those  started  on  XI, 
13,  1911,  allowing  for  the  slight  and  decreasing  differences  due  to  age,  and 
of  both  sets  of  cultures  of  Gymnogramme  triangularis.  How  can  one  account 
for  these  differences?  The  cultures  were  sowed  all  together,  the  culture  solu- 
tion, the  tiles  and  the  dishes  had  all  been  treated  exactly  alike  and  together; 
no  selections  were  made  at  any  time.  Some  of  the  cultures  were  put  on  the 
turn-tables  at  once  after  sowing  and  the  remainder  were  placed  on  the  shelf  by 
them.  They  were  thereupon  marked.  Almost  from  the  moment  when  the 
dry  spores  touched  the  moist  tile  they  began,  either  to  remain  still,  or  to  turn, 
with  the  tile  on  which  they  had  fallen.  And  so  they  remained  night  and  day, 
the  diffused  sunlight  from  the  window  falling  through  a  white  Holland 
shade  nearly  horizontally  upon  them  by  day,  darkness  enveloping  them  at 
night  (for  I  very  seldom  use  artificial  light  in  the  room  where  the  multiple 
clinostat  is),  turning  night  and  day  or  staying  motionless,  according  to  their 
position;  watered  from  time  to  time  with  fresh  Knopp's  solution  when  nec- 
essary, equally  warmed  and  similarly  treated  in  every  respect,  so  far  as  I 
can  see,  except  that  in  one  respect  they  are  not  similarly  lighted.  The  light 
is  the  same  in  composition,  intensity  and  duration,  not  in  direction.  This  dif- 
ference alone  is  accompanied  by  the  differences  in  the  vegetative  and  the  re- 
productive parts  above  described. 

It  may  be  easier  to  gain  some  insight  into  this  problem,  into  the  reasons 
or  causes  of  these  differences,  if  we  consider  the  vegetative  and  the  repro- 
ductive parts  separately.  Acting  on  this  principle,  I  proceeded  to  experiment 
upon  young  flowering  plants  grown  from  the  seed. 

SEEDLINGS  OF   MUSTARD  AND  OF   WHEAT. 

Seedlings  of  white  mustard  (Sinapis  alba)  and  of  wheat  were  sowed  on 
sterilized  greenhouse  soil  in  2-inch  porous  flower  pots.  I  selected  these 
plants  because  of  the  promptness  with  which  they  germinate  and  the  vigor 
with  which  they  grow,  for  a  time  at  least,  under  laboratory  conditions,  and 
because  the  early  growth  of  the  one  (mustard)  is  mainly  hypocotyledonary 
and  at  the  expense  of  food  made  by  the  seedling  as  well  as  drawn  from  the 
seed,  whereas  the  growth  of  the  other  (wheat)  is  mainly  epicotyledonary  and 
the  seedling,  though  well  fed,  is  not  self-nourishing  for  some  time.  Recalling 
the  well  known  phototropism  of  these  two  seedlings,  I  thought  that  by  ex- 
posing the  two  sets,  one  on  the  shelf  and  one  on  the  turn-tables,  to  the  same 
light,  I  could  ascertain  whether  there  were  any  greater  stimulus  to  growth 
for  the  one  set  of  plants  or  the  other,  whether  if  an  adjustment  as  to  position 
between  light  and  darkness — that  is,  between  more  and  less  light — cannot  be 
attained,  growth  will  be  more  rapid  than  where  a  plant  is  able  to  attain  a 


STUDIES    OF    IRRITABILITY    OF    PLANTS PEIRCE  75 

position  of  such  adjustment.  A  geranium,  for  example,  growing  on  a  window 
sill,  turns  toward  the  light.  If,  after  it  has  accomplished  a  bend  toward 
the  window,  it  be  turned  around,  it  will  reverse  the  bend  or  make  a  new  one, 
again  carrying  the  tip  over  toward  the  light.  And  this  process  may  be 
repeated  indefinitely,  with  the  same  result  so  long  as  the  plant  can  grow. 
Such  a  plant  is  likely  to  become  longer  in  the  same  length  of  time  than  one 
beside  it  which  has  not  been  changed  in  position.  If  this  is  the  effect  on 
stems,  the  position  of  which  is  reversed  only  at  long  intervals — say  every 
other  day  or  two — would  this  also  be  the  case  if  the  intervals  were  short? 
The  result  of  an  experiment  on  mustard  will  throw  some  light  on  this  question. 
Seeds  of  Sinapis  alba  were  sowed  on  II,  22,  1912,  on  greenhouse  soil  in 
2-inch  flower  pots,  six  of  which  were  put  on  turn-tables  making  four  revolu- 
tions a  minute,  seven  on  turn-tables  making  two  revolutions  each  minute,  and 
seven  on  the  shelf,  and  therefore  getting  light  mainly  from  one  side.  These 
were  allowed  to  grow  until  the  first  leaves  in  the  plumule  began  to  show 
and  were  thereupon  measured,  that  is,  on  III,  12,  1912,  nineteen  days  after 
sowing.  The  length  taken  for  measurement  was  that  from  the  surface  of 
the  soil  to  the  tip  of  the  plant.  The  data  follow : 

Average  length  of  29  seedlings  in  Pot    I  on  M-minute  turn-table.  4.08  cm. 
16  "  "        II  3.88 

13  "  "      III  4.13 

9  "  "       IV  4.53 

29  "  "        V  4.96 

5  "  «      VI  4.38 


101  4.32  " 

Average  length  of  24  seedlings  in  Pot    I  on  3^-minute  turn-table.  3.90  cm. 

24  "  "  II  4.25 

19  "  "  III  3.92 
10  "  "  IV  3.20 

25  "  "  V  4.78 
23  "  «  VI  4.01 

6  "  "  VII  4.09 

131  "  4.02  " 

Average  length  of     6  seedlings  in  Pot     I  on  the  shelf .  4.27cm. 

15  "  "  II  4.80 

20  "  "  III  3.80 
12  "  "  IV  3.99 
15  V  4.89 
19  "  "  VI  4.33 
14  "  "  VII  3.91 

101  "  4.28  " 


76  DUDLEY    MEMORIAL   VOLUME 

From  these  three  averages  of  the  lengths  of  mustard  seedlings — 4.32 
cm.,  4.02  cm.  and  4.28  cm. — it  is  clear  that,  so  far  as  the  growth  of  the 
hypocotyledonary  stem  is  concerned,  it  makes  no  material  difference  whether 
the  plant  is  illuminated  mainly  from  one  side  or  on  all  sides  successively.  So 
far  as  I  could  see,  the  seedlings  all  presented  a  normal  appearance,  both  in 
stems  and  cotyledons,  as  to  size,  color  and  form. 

This  experiment  having  failed  to  throw  any  light  on  the  question,  I 
sowed  wheat  similarly  on  III,  15,  1912,  and  put  five  pots  on  the  quarter- 
minute  turn-tables,  five  pots  on  the  half-minute  turn-tables  and  five  on  the 
shelf  beside  them.  Four  weeks  after  sowing  I  measured  them,  from  the  sur- 
face of  the  soil  to  the  tip  of  the  unopened  leaf,  with  the  following  results : 

Average  length  of  10  seedlings  in  Pot    I  on  M-minute  turn-table.  13.2  cm. 
7          "  "       II  11.6 

7          "  "      III  15.2 

3  "IV  16.2 
7           «           «        V                                             17.3 

34  "  14.7     " 

Average  length  of    9  seedlings  in  Pot    I  on  H-minute  turn- table.  17. 1  cm. 

10  "  "II  16.3 
12           "            "      III  9.6 

11  "  "       IV  12.9 

4  "V  14.0 

«  «          46  "  13.98  " 

Average  length  of    6  seedlings  in  Pot    I  on  the  shelf.  9 . 68  cm. 

9  "  "II  18.63 

10  "  "      III  13.45 

6  "  "IV  12.7 

10  "  "        V  15.3 


41  13.95  " 

This  experiment  also,  so  far  as  the  growth  of  the  stem  is  concerned, 
throws  no  light  on  our  question,  for  there  is  no  material  difference  in  these 
averages,  of  14.7  cm.,  13.98  cm.  and  13.95  cm.  There  was,  however,  a 
difference  in  the  length  of  the  leaves,  which  unfortunately  I  did  not  realize 
in  time  to  measure.  But  the  leaves  of  the  seedlings  on  the  turn-tables  were 
in  many  instances  as  long  again  as  those  of  seedlings  on  the  shelf.  Here, 
then,  we  do  have  a  resemblance  to  the  behavior  of  the  f  ern-prothallus  and  the 
liverworts.  In  both  instances  we  have  chlorophyll-containing  photosyntheti- 


STUDIES    OF    IRRITABILITY    OF    PLANTS PEIRCE  77 

cally  food-manufacturing  organs,  dependent  for  their  efficiency  upon  the 
amounts  of  light  and  of  carbon  dioxide  which  can  penetrate  to  the  deeper  as 
well  as  more  .superficial  cells.  I  cannot  see  that  the  revolution  of  a  culture 
in  a  covered  crystallizing  dish  at  no  greater  speed  than  four  times  a  minute 
could  promote  diffusion  or  otherwise  increase  the  supply  of  carbon  dioxide 
sufficiently  to  account  for  the  larger  size  of  the  prothalli  and  of  the  liver- 
worts on  the  turn-tables,  as  compared  with  those  motionless  on  the  shelf. 
The  supply  of  carbon  dioxide  to  the  leaves  of  wheat  seedlings  revolving  on 
turn-tables  may  be  greater  than  for  the  plants  stationary  on  the  shelf.  But 
the  supply  of  light  is  certainly  increased  in  the  same  way  that  the  supply  of 
heat  is  increased  for  the  man  who  first  turns  his  face  and  afterwards  his  back 
to  a  fire  until  he  is  comfortably  warmed. 

DISCUSSION    AND    SUMMARY. 

1.  An  analysis  of  the  influence  of  light  upon  growth  in  plants  shows 
that  it  affects  the  direction,  kind,  rate  and  amount  of  growth12.  Phototropic 
bending  of  plants  and  plant-organs  is  of  common  and  long-established 
knowledge.  That  the  kind  of  growth  is  influenced  by  light  has  been  shown 
mainly  by  the  earlier  ecologists,  such  as  Stahl13,  and  by  the  experi- 
mental morphologists  and  physiologists  like  Goebel14,  Klebs15,  and 
Vochting16.  It  has  long  been  supposed  that  the  rate  of  growth,  and  also 
the  amount,  are  greatly  influenced  by  light,  that,  as  Sachs17  would  have  it, 
light  depresses  the  rate  of  growth,  other  things  being  equal.  Common  ex- 
perience shows  that  plants  grown  in  darkness  or  in  insufficient  light  are  long, 
slender,  spindling,  or  at  least  "drawn,"  as  compared  with  plants  growing  for 
the  same  length  of  time,  and  under  otherwise  identical  conditions,  in  the 
light.  But  that  the  cause  of  this  difference  can  be  expressed  in  the  usual 


«  Peirce,  G.  J.    Text  book  of  Plant  Physiology,  p.  210.  1903. 

13  Stahl,  F.     Ueber  den  Einfluss  des  sonnigen  oder  schattigen  Standortes  auf 
die  Ausbildung  der  Laubblatter.     Zeitschr.  f.  Naturwissenschaft,  XVI.     Review  in 
Botanische  Zeitung,  41,   1883. 

14  Goebel,  K.     Einleitung  in  die  experimentelle  Morphologic  der  Pflanzen.     1908. 
And  the  literature  there  cited. 

15  Klebs,  G.     Willkurliche  Entwickelungsanderungen  bei  Pflanzen.     1903.     And 
the  literature  there  cited. 

16  Vochting,  H.     Ueber  den  Einfluss  des  Lichtes  auf  die  Gestaltung  und  Anlage 
der  Bliiten.    Jahrb.  f.  wiss.  Bott.  XXV.     1893. 

17  Sachs,  J.     Lectures  on  the  Physiology  of  Plants.     Oxford,  1887.     Etc. 


78  DUDLEY    MEMORIAL    VOLUME 

simple  terms  I  believe  the  experiments  which  are  described  in  the  preceding 
pages  show  is  not  the  case.  Light  does  not  always  check  growth,  other  things 
being  equal.  Instead,  it  promotes  it,  also,  other  things  being  equal. 

When  plants  are  grown  in  darkness,  their  leaves  are  small  and  may  even 
be  misshapen ;  it  is  their  stems  which  are  long,  and  they  lack  thickness.  The 
organs  devoted  to  food  manufacture  require  light  for  their  normal  develop- 
ment. This,  too,  has  long  been  known  and  it  has  been  explained  on  the 
supposition  that  use  (activity)  and  food  are  necessary  for  the  development 
of  leaves  and  their  tissues.  Given  a  certain  amount  of  light,  growth  (and 
development)  should  take  place  at  a  certain  rate  and  to  a  certain  amount, 
other  things  being  equal.  By  a  certain  amount  of  light  is  meant  that  quan- 
tity which  the  plant  or  organ  can  use  or  which  so  penetrates  its  living  cells 
as  to  affect  them.  We  conceive  that,  under  usual  conditions,  a  plant  and  its 
organs  grow  in  direction,  rate  and  amount  into  such  positions  as  afford  what 
may  be  called  the  optimum,  all  its  activities  contributing  to  this  resultant.  If 
this  be  true,  the  position  and  size  of  a  plant  represent  the  influence  of  cir- 
cumstances upon  its  substance.  By  modifying  the  circumstances  in  any 
way,  we  may  also  influence  the  resultant.  Without  increasing  the  quantity 
of  light  available  or  changing  the  quality,  and  without,  so  far  as  I  can  see, 
affecting  the  supply  of  carbon  dioxide  and  of  other  food  materials,  the  plants 
used  in  the  experiments  here  described  exhibit  remarkable  differences  accord- 
ing to  their  different  exposures  to  light.  The  usual  position  occupied  in  the 
light  by  chlorophyll-containing  organs  is  that  which  presents  the  greatest 
possible  contrast  between  the  amounts  of  light  available  on  the  two  sides  of 
these  organs.  The  positions  of  ordinary  leaves  show  this,  and  I  am  not  sure 
that  even  such  vertical  and  two-faced  leaves  as  those  of  Eucalyptus  do  not  also 
show  this  more  or  less.  But  in  darkness  and  on  turn-table  there  is  no  such 
position  of  contrast.  There  is  no  contrast.  In  darkness — that  is,  under  con- 
ditions free  from  the  alleged  depressing  influence  of  light  upon  growth — 
growth  does  not  take  place  to  the  usual  extent  in  chlorophyll-containing  or- 
gans, even  when  food  is  supplied  in  adequate  quantities  and  in  suitable  form. 
Under  these  conditions  growth  should  be  greater.  But  one  may  attribute  this 
lesser  growth  to  lack  of  use  of  the  chlorophyll  apparatus.  This  hypothesis 
may  usually  be  correct  without  necessarily  being  complete.  For  light  may  con- 
ceivably stimulate,  apart  from  its  effect  on  food  manufacture  and  the  appa- 
ratus concerned.  The  manufacture  of  food  depends  upon  many  factors,  of 
which  sufficient  water,  light,  carbon  dioxide  and  chlorophyll  are  the  most 


STUDIES    OF    IRRITABILITY    OF    PLANTS PEIRCE  79 

obvious.  The  available  amounts  of  these  decrease  in  the  order  named,  the 
supply  of  chlorophyll  pigments  being  always  very  small  but  also  very  effect- 
ive. The  supply  of  carbon  dioxide,  because  of  its  great  dilution,  rarely 
reaches  the  optimum  proportion,  whereas  the  amounts  of  chlorophyll,  light 
and  water,  in  proportion  to  carbon  dioxide,  often  exceed  it.  Increasing  the 
supply  of  light  beyond  the  usual  amount  without  a  corresponding  increase 
in  the  supply  of  carbon  dioxide  does  not  necessarily  increase  the  products  of 
photosynthesis.  But  the  foregoing  experiments  show  that  increased  exposure 
of  chlorophyll-containing  organs  of  liverworts,  ferns  and  one  grass  to  the 
usual  quantity  of  light  results  in  increased  growth.  This  increased  growth 
necessarily  implies  increased  use  of  food,  perhaps  there  may  also  be  increased 
food  manufacture,  but  I  do  not  yet  know  this  to  be  the  case.  From  this, 
one  is  forced,  so  far  as  I  can  see,  to  conclude  that  light,  up  to  a  certain  in- 
tensity at  least,  stimulates  growth  rather  than  depresses  it. 

2.  We  have  seen,  also,  that  a  more  uniform  illumination  increases  sym- 
metry in  development.     In  certain  instances  this  symmetry  appears  to  be 
complete,  a  dorsi-ventral   giving  place  to  radial  structure,   the  stimulating 
effect  of  light  showing  itself,  not  merely  in  change  (increase)  in  size,  but  also 
in  change  in  form. 

3.  The  foregoing  experiments  on  the  ferns,  Pteris  aquilina  and  Gym- 
no  gramme  triangularis,  show  that,  although  the  archegonia  and  antheridia 
ordinarily  form  on  the  shaded  side  of  the  prothallus,  they  form  on  both  sides 
when  the  illumination  is  equal,  and  in  greater  numbers  on  the  two  sides  than 
is  usual  on  either.     Increased  illumination  is  followed  and  accompanied  by 
increased  numbers  of  the  organs  of  sexual  reproduction,  the  fertility  of  which 
is  evident  from  the  development  of  sporophytes  whenever  there  is  sufficient 
water  for  the  sperms  to  swim.     I  am  not  aware  that  such  experiments  have 
been  made  on  ferns,  but  the  experiments  of  Klebs18,  Vochting19  and  their 
followers  on  algae,  fungi  and  the  flowering  plants,  demonstrate  the  intimate 
connection  between  illumination  and  the  development  of  reproductive  organs, 
light  being  for  many  plants  the  indispensable  stimulus  thereto.     The  behavior 
of  ferns  is,  therefore,  consistent  with  the  behavior  of  the  other  plants  already 
experimentally  investigated. 


18  Klebs,  G.    Die  Bedingungen  der  Fortpflanzung  bei  einigen  Algen  und  Pilzen. 
Jena.     18%. 

19  Vochting,  H.    Ueber  den  Einfluss  des  Lichtes  auf  die  Gestaltung  und  Anlage 
der  Bliiten.    Jahrb.  f.  wiss.  Bott.  XXV.     1893. 


80  DUDLEY    MEMORIAL   VOLUME 

4.  As  to  the  possible  influence  of  light  on  the  ratio  of  male  and  female 
prothalli,  I  wish  merely  to  record  the  observation  that,  in  my  cultures,  illumi- 
nation, increased  by  revolving  the  prothalli,  seemed  to  be  followed  by  an 
increased  proportion  of  female  prothalli.  I  do  not  care  at  this  time  to  dis- 
cuss this  observation  or  to  hazard  an  opinion  as  to  the  extent  of  this  possible 
effect  of  light  on  plants. 


Fig.   1.     Fern  prothalli   five  months  old,  grown 

from   spores  on   a   turn-table   revolving   four 

times  a  minute. 
Fig.   3.     Fern  prothalli  of  same  age,  from  same 

lot  of  spores,   receiving  light   from  one  side 

only. 


Fern   prothalli   five  months  old,  grown 
spores  on   a   turn-table   revolving   four 
times  a  minute. 
Fig.   4.     Fern  prothalli  of  same  age,  from  same 
lot  of  spores,   receiving  light   from  one  side 
only. 


THE     GYMNOSPERMS     GROWING     ON     THE     GROUNDS     OF 
LELAND     STANFORD     JR.     UNIVERSITY. 

LERov  ABRAMS,  Associate  Professor  of  Botany. 

THE   GYMNOSPERMS   are  biologically  more   primitive  than  the  Angio- 
sperms.     The  flowers  are  always  unisexual  and  without  perianth  (ex- 
cept Gnetales1).     The  staminate  flowers  resemble  those  of  the  club- 
mosses  and  are  short  or  elongated  shoots  bearing  a  number  of  spiral  or  virti- 
cillate  stamens.     The  ovulate  flowers  are  of  a  more  varied  structure,  but  the 
ovules  are  not  enclosed  in  an  ovary  as  in  the  Angiosperms. 

The  Gymnosperms  comprise  only  trees  and  shrubs,  and  are  represented 
by  four  living  and  two  extinct  orders.  Representatives  of  two  of  the  orders, 
Ginkgoales  and  Conijerce,  are  cultivated  on  the  University  grounds.* 

GINKGOALES. 

A  single  species,  Ginkgo  biloba,  is  the  sole  survivor  of  this  ancient  order 
of  Gymnosperms. 

1.    Ginkgo  biloba  L.     Ginkgo  or  Maidenhair  Tree. 

Leaves  deciduous,  clustered  on  short  stubby  twigs,  fan-shaped,  thickened 
on  the  margin  and  usually  divided,  parallel-veined;  flowers  dioecious;  stami- 
nate in  slender  aments ;  ovulate  in  pairs  on  long  stalks ;  fruit  drupe-like,  with 
an  ill-scented,  fleshy  coat  surrounding  a  smooth  oval  stone. 

Native  of  China  and  Japan.  Handsome  staminate  trees  are  on  the 
grounds  of  the  Stanford  Residence,  and  a  few  young  trees  are  planted  on  the 
Campus. 

CONIFERS. 

Trees  or  shrubs  with  more  or  less  resinous  wood  and  usually  narrowly 
linear  or  needle-like  evergreen  or  rarely  deciduous  leaves.  Flowers  monoe- 
cious or  dioecious.  Fruit  a  woody  cone  or  fleshy  and  drupe-like. 


*  In  the  spring  of  1909  Professor  Dudley  prepared  a  key  to  the  Conifers  grow- 
ing on  the  Campus  for  the  use  of  the  students  of  Forest  Botany.  As  only  a  few 
carbon  copies  were  made  the  present  paper  was  undertaken  in  order  that  Pro- 
fessor Dudley's  work  might  be  embodied  in  permanent  form.  Although  the  key 
ha.s  been  the  basis  of  this  paper  the  writer  has  gone  over  the  field  with  considerable 
thoroughness,  verifying  the  identifications,  and  has  added  the  following  species  not 
credited  to  the  Campus  by  Professor  Dudley:  Agathis  loranthifolia,  Cephalotaxus 
pedunculata,  Picea  Parryana,  P.  sitchensis,  Pinus  Cembra,  P.  edulis,  P.  excelsa, 
P.  Jeffreyi,  P.  Lambertiana,  P.  monticola,  P.  nigra,  P.  Pinea,  Sciadopitys  verticillata 
(probably  added  to  our  collections  since  the  key  was  prepared),  Taxodium  mucrona- 
tum,  Thuyopsis  dolabrata. 


82  DUDLEY    MEMORIAL   VOLUME 

The  Coniferse  are  the  prevailing  Gymnosperms  of  the  present  geological 
age.     They  are  represented  by  two  families  and  about  forty  genera. 
Fruit  composed  of  a  solitary  stone  surrounded  by  a  fleshy,  greenish  or  bright 

red  aril.  1.  Taxacea. 

Fruit  a  woody  cone  made  up  of  scales,  each  of  which  bears  1 -several  seeds  in 

its   axil    (scales  coalescent   and  fleshy   in  Juniperus). 

2.  Pinacea. 

1.    TAXACE^E.     Yew  Family. 

Trees  or  shrubs  with  durable  close-grained  wood.  Leaves  flat,  linear 
or  broader,  usually  2 -ranked  by  a  twist  of  the  compressed  petioles,  decur- 
rent  on  the  branchlets.  Flowers  dioecious,  axillary;  staminate  composed  of 
many  stamens;  ovulate  of  a  single  erect  ovule.  Fruit  a  stone  more  or  less 
enclosed  by  a  fleshy  aril-like  disk. 
Fruit  drupe-like;  aril  completely  enclosing  stone. 

Leaves  broadly  lanceolate;  pollen-sacs  2.  1.  Podocarpus. 

Leaves  linear  or  nearly  so;  pollen-sacs  3-4. 

Flowers  pedunculate,   clustered  in  heads;   pollen-sacs  3. 

2.  Cephalotaxus. 
Flowers  axillary;   pollen-sacs  4.  3.  Torreya. 

Fruit  with  a  bright  red  cup-shaped  aril  partly  enclosing  the  stone. 

4.   Taxus. 

1.  Podocarpus. 

Trees  or  sometimes  shrubs  with  linear  or  lanceolate  alternate  or  oppo- 
site evergreen  leaves.  Fruit  drupe-like,  borne  on  a  thickened  foot-stalk; 
seeds  inverted. 

A  genus  of  over  40  species,  chiefly  in  the  tropics  and  substropices  of  the 
southern  hemisphere.  Many  species  are  valuable  timber  trees. 

1.  Podocarpus  macrophylla  Don. 

Leaves  alternate,  lanceolate,  sometimes  falcate,  2-5  inch  long,  about 
Y-2.  inch  wide,  bright  green  and  lustrous  above,  pale  beneath;  fruit  greenish, 
on  a  thickened  purplish  foot  stalk. 

One  specimen,  planted  by  Professor  Dudley,  is  on  the  lawn  at  8  Al- 
varado  Row. 

2.  Cephalotaxus. 

Leaves  linear,  acute  and  often  sharp-pointed,  spirally  arranged  but  usu- 
ally appearing  2-ranked.  Flowers  dioecious,  pedunculate,  clustered  in  small 
heads.  Fruit  drupe-like ;  endosperm  uniform. 


GYMNOSPERMS   ON    STANFORD   GROUNDS ABRAMS  83 

A  small  genus  containing  but  4  species,  all  of  which  are  native  of  Japan 
and  China. 
Leaves  appearing  2-ranked  by  a  twist  in  the  petiole. 

Leaves  y-z'Y^  incn  l°ng>  abruptly  acute.  1.  C.  drupacea. 

Leaves  about  2  inches  long,  gradually  tapering  from  near  base. 

2.  C.  Fortunei. 
Leaves  spreading  from  all  sides,  at  least  on  the  principal  shoots,  spiny-tipped. 

3.  C.  pedunculata. 

1.  Cephalotaxus  drupacea  Siebold. 

Leaves  appearing  2-ranked  and  opposite,  l/2-^.  inch  long,  scarcely  j/£ 
inch  wide,  abruptly  acute,  margins  not  revolute;  fruit  elliptic,  about  ^4 
inch  long ;  stone  smooth. 

Native  of  northern  China  and  Japan,  where  it  attains  a  height  of  30-40 
feet.  One  ovulate  shrub  is  at  the  west  end  of  Encina  Garden  and  a  couple 
of  small  staminate  specimens  are  in  the  nursery. 

2.  Cephalotaxus  Fortunei  Hook. 

Leaves  2-ranked,  margins  thin,  slightly  revolute,  gradually  tapering  to 
the  sharp-pointed  apex,  about  2  inches  long,  over  l/%  inch  wide. 

Tree  40-60  feet  high  with  long,  slender,  drooping  branches.  Native  of 
northern  China.  One  small  staminate  tree  is  north  of  the  live  oak  on  the 
Mausoleum  lawn. 

3.  Cephalotaxus  pedunculata  Siebold. 

Leaves  on  leading  shoots  and  principal  branches  scattered  on  all  sides 
of  the  stem,  \y2  inches  long,  sharply  acute,  margins  distinctly  revolute. 

A  small  Japanese  tree,  20-25  feet  high,  with  the  spreading  branches  in 
whorls. 

One  small  sterile  shrub  in  very  poor  condition  is  in  the  eastern  part  of 
the  Cactus  Garden. 

3.  Torreya. 

Leaves  flat  and  linear,  appearing  2-ranked,  spiny-tipped.  Flowers 
dioecious,  axillary;  pollen  sacs  4,  arranged  in  a  semi-circle;  fruit  drupe-like, 
the  greenish  aril  completely  enclosing  the  stone ;  endosperm  nutmeg-like. 

A  genus  of  four  species,  widely  separated  geographically  and  of  very 
local  distribution.  One  is  in  Florida,  one  in  California,  and  the  other  two 
are  in  China  and  Japan. 


84  DUDLEY    MEMORIAL   VOLUME 

1.    Torreya  calif ornica  Torrey.     California  Nutmeg. 

Leaves  \l/2-l  inches  long,  %  inch  wide,  tapering  slightly  to  the  sharp- 
pointed  apex,  dark  glossy  green  above,  pale  beneath;  fruit  \-ll/2  inches  long. 

Tree  sometimes  80  feet  high,  with  spreading  or  drooping  branches  and 
smooth,  scaly  bark.  Native  of  central  and  northern  California  in  the  Coast 
Ranges  and  the  Sierra  Nevada. 

4.  Taxus.     Yew. 

Trees  or  shrubs  with  spreading  or  erect  branches  and  scaly  bark. 
Leaves  flat,  linear,  2-ranked.  Flowers  dioecious,  axillary;  pollen-sacs  6-8, 
arranged  in  a  circle.  Fruit  with  a  bright  red  fleshy  viscid  open  cup  partly 
enclosing  the  erect  stone.  • 

A  genus  of  approximately  8  closely  related  species,  distributed  through 
the  north  temperate  regions. 

1.  Taxus  baccata  L.     English  Yew. 

Leaves  linear,  2-ranked,  usually  falcate,  shortly  accumulate,  dark  green 
above,  pale  beneath,  l/4-fy  inch  l°ng>  iruit  l/3-l/2  inch  broad,  with  almost 
globose  aril. 

The  English  Yew  is  a  native  of  Europe,  western  Asia  and  northern 
Africa.  It  has  long  been  cultivated  and  many  garden  forms  exist.  Several 
shrubs  are  in  the  neighborhood  of  the  Cactus  Garden,  especially  near  the 
entrance. 

la.  Taxus  baccata  fastigiata  Loud.     Irish  Yew. 

Leaves  linear,  as  in  the  typical  form,  but  spreading  from  all  sides  of  the 
erect  fastigiate  branches. 

Specimens  are  on  the  borders  of  the  Cactus  Garden. 

Ib.  Taxus  baccata  adpressa  Carr. 

This  variety  is  distinguished  by  its  broader  and  shorter,  oblong  obtuse 
leaves. 

Our  specimens  are  the  compact  erect  form^  They  are  on  the  east  and 
west  sides  of  the  Cactus  Garden. 

2.  PINACE.E.     Pine    Family. 

Trees  or  shrubs  with  resinous  wood.  Leaves  linear,  needle-like  or 
scale-like,  spirally  arranged  or  decussately  opposite,  evergreen  or  in  a  few 
genera  deciduous.  Flowers  dwecious;  staminate  in  ament-like  deciduous 
clusters;  pollen-sacs  2-several.  Fruit,  a  woody  cone,  made  up  of  several 
scales,  each  with  or  without  a  subtending  bract,  bearing  1  or  more  seeds  at 


GYMNOSPERMS   ON    STANFORD   GROUNDS ABRAMS  85 

base  on  the  upper  surface  (scales  coalescent  and  fleshy  in  Juniperus]  ;  seeds 

often  winged. 

Leaves  and  cone-scales  spirally  arranged. 

Leaves  usually  broad  at  base  and  tapering  to  a  sharp  point  ;  ovules  and 

seed  1  to  each  scale.  Tribe  1.  Araucarece. 

Leaves  linear  or  needle-like. 

Leaf -bases  not  decurrent  on  twigs ;  ovules  and  seeds  2  to  each  scale, 

pendent.  Tribe  2.  AbietecB. 

Leaf-bases  decurrent  on  twigs;  ovules  and  seeds    several  to  each  scale, 

erect.  Tribe  3.  Taxodece. 

Leaves  and  cone- scales  opposite,  the  former  often  scale-like. 

Tribe  4.  Cupressece 
Tribe   1.     Araucareae. 

A  very  distinct  tribe  composed  of  the  two  genera  A  gat  his  and  Araucaria. 
These  are  almost  wholly  restricted  in  their  distribution  to  the  tropical  or 
extra-tropical  realms. 

Scales  persistent;  seeds  free.  1.  Agathis. 

Scales  deciduous;  seeds  adherent.  2.  Araucaria. 

1.  Agathis. 

Leaves  opposite  or  alternate,  ovate-lanceolate,  attenuate,  parallel-veined 
and  of  a  firm  leathery  texture.  Flowers  dioecious,  solitary,  the  staminate 
elongated.  Cones  ovate  to  globose,  axillary;  scales  persistent;  seeds  winged 
only  on  one  side,  free. 

1.    Agathis  loranthifolia  Salisbury.     Amboyna  Pine. 

Leaves  opposite  or  sometimes  alternate,  1^-4  inches  long,  ovate-lanceo- 
late, glaucous  green;  staminate  flowers  2  inches  long;  cones  globose  or  tur- 
binate,  3-4  inches  long. 

A  large  forest  tree,  often  8  to  10  feet  in  diameter  and  100  feet  or  more 
high.  Native  of  the  mountains  of  the  Malay  Archipelago.  One  young 
specimen,  which  seems  perfectly  hardy  in  this  climate,  is  in  Professor 
Durand's  garden. 

2.  Araucaria. 

Leaves  evergreen,  flattened,  lanceolate  or  short-ovate  to  awl-shaped, 
usually  sharp-pointed.  Flowers  normally  dioecious ;  cones  erect,  their  scales 
deciduous;  ovules  and  seeds  1  to  each  scale,  pendent;  anthers  with  several 
elongated  pendent  pollen-sacs. 

Araucaria  is  represented  by  10  species,  all  of  which  are  restricted  to  the 
southern  hemisphere. 


86  DUDLEY    MEMORIAL    VOLUME 

Leaves  lanceolate  to  ovate. 

Leaves  lanceolate;  spiny  tip  of  scale  about  ^  inch  long. 

Leaves  2-ranked,  dark  green;  scales  nearly  as  broad  as  long,  bract  and 

scale  distinct  toward  the  apex.  1.  A.  Bidwillii. 

Leaves  spreading  from  all  sides  of  the  branches;  scales  over  twice  as 
long  as  broad,  bract  and  scale  completely  united. 

2.  A.  braziliana. 
Leaves  broadly  ovate,  concave,  stiff  and  very  spiny-tipped;  spiny  tip  of 

scale  \l/2  inches  long.  3.  A.  imbricata. 

Leaves  awl-shaped;  bark  exfoliating.  4.  A.  Cunninghamii. 

1.  Araucaria  Bidwillii  Hooker. 

Leaves  narrowly  ovate-lanceolate,  flat,  rigid,  sharp-pointed,  dark  green 
and  shiny,  those  on  the  branches  twisted  and  2-ranked,  1-2  inches  long;  cones 
ovate-globose,  8-10  inches  long;  scale  and  bract  distinct  above  the  middle, 
free  part  of  scale  fy  inch  broad;  bract  2  inches  long,  nearly  as  wide,  thin, 
sharply  and  very  prominently  keeled  at  apex,  terminated  by  a  reflexed  linear- 
lanceolate  spine,  24  inch  long. 

A  forest  tree  100  to  150  feet  high.  Native  of  the  Brisbane  Mountains, 
Australia,  where  it  is  known  as  the  Bunya-Bunya.  A  large  tree  is  on  the 
grounds  of  the  Stanford  Residence,  and  smaller  specimens  are  near  the 
Mausoleum  Avenue  and  in  Encina  Garden. 

2.  Araucaria  braziliana  Rich. 

Leaves  linear-lanceolate,  tapering  to  a  spiny  tip,  flat,  straight,  spreading 
from  all  sides  of  the  branches,  rather  pale  glaucous  green,  1-2  inches  long; 
cones  globose,  6-8  inches  long;  bracts  and  scales  completely  united  and  in- 
distinguishable, wedge-shaped  and  somewhat  4-sided,  2  inches  long,  Y^  inch 
broad,  terminated  by  a  linear-lanceolate  spine,  J^  inch  long. 

A  tree  75  to  100  feet  high.  Native  of  southern  Brazil.  Two  small 
trees  are  between  the  Cactus  Garden  and  the  Mausoleum. 

3.  Araucaria  imbricata  Pavon.     Monkey  Puzzle. 

Leaves  broadly  ovate,  closely  set  and  spreading  from  all  sides  of  the 
branches,  straight,  concave,  rigid,  very  sharp-pointed,  £4-1  inch  long;  cones 
globose,  6-8  inches  long;  scales  wedge-shaped,  2  inches  long,  about  1  inch 
broad,  terminated  by  a  slender  subulate  spine  1^2  inches  long. 

A  forest  tree,  often  100  feet  high,  forming  extensive  forests  in  the 
Andes  of  southern  Chili.  A  good  specimen  is  on  the  Mausoleum  lawn,  and 
other  smaller  trees  are  on  the  Campus. 


GYMNOSPERMS   ON    STANFORD   GROUNDS ABRAMS  87 

4.  Araucaria  Cunninghamii  Ait. 

Leaves  awl-shaped,  enlarged  at  base,  yz  inch  long,  spreading  from  all 
sides  of  the  branches,  rigid ;  cones  ovate-globose,  3-4  inches  long ;  scales 
wedge-shaped,  ^  inch  broad,  terminated  by  an  awl-shaped  awn  l/$  its  length. 

A  large  tree,  100  to  150  feet  high,  with  exfoliating  bark.  An  Australian 
species  forming  extensive  forests  in  New  South  Wales  and  Queensland.  One 
tree  is  at  the  east  end  of  Roble  Garden  and  another  is  on  the  southwest 
border  of  the  Cactus  Garden. 

Araucaria  excelsa  R.  Br.  The  Norfolk  Island  Pine  resembles  A.  Cun- 
ninghamii, but  the  branches  are  in  very  symmetrical  and  rather  distant  whorls. 
This  species  is  not  cultivated  on  the  grounds,  but  is  planted  in  gardens  at 
San  Jose  and  San  Francisco. 

Tribe    2.     Abieteae. 

Leaves  and  floral  parts  spirally  arranged;  ovulate  scales  subtended  by 
bracts,  becoming  woody  and  forming  a  cone  in  fruit;  ovules  2,  adnate  to 
the  upper  surface  of  scale  near  the  base,  pendent ;  seeds  usually  with  a  con- 
spicuous membranous  wing;  cotyledones  3  or  more. 
Leaves  in  1-5-leaved  clusters,  surrounded  at  base  by  membranous  sheaths; 

cones  maturing  the  second  year.  1.  Pinus. 

Leaves  without  basal  sheaths,  scattered  or  clustered  on  short,  stubby  branch- 
lets  ;  cones  maturing  the  first  year.          » 
Leaves  clustered  on  short,  stubby  branchlets;  cones  erect. 

2.  Cedrus. 

Leaves  single,  spirally  arranged  or  appearing  2-ranked. 
Cones  pendent ;  scales  persistent. 

Branchlets  rough  with  the  persistent,  woody  leaf-bases ;  bracts  not 

exserted.  3.  Picea. 

Branchlets  smooth ;  leaf -bases  sessile ;  bracts  3-parted,  well  exserted. 

4.  Pseudotsuga. 
Cones  erect;  scales  deciduous;  branchlets  smooth. 

5.  Abies. 
1.  Pinus.     Pines. 

Trees  or  a  few  shrubs.  Leaves  evergreen,  needle-shaped,  in  clusters  of 
2-5  (solitary  in  one  species)  from  the  axils  of  scale-like  primary  leaves,  each 
cluster  surrounded  at  base  by  a  persistent  or  deciduous  sheath  of  membra- 
nous scales;  cones  maturing  the  second  year,  their  scales  persistent,  woody, 
often  thickened  or  awned  with  a  prickle  at  apex.  Seeds  usually  winged. 

The  pines  with  approximately  eighty  living  species  constitute  the  largest 
genus  of  the  Gymnosperms.  They  are  restricted  to  the  northern  hemisphere 
and  chiefly  to  the  temperate  regions. 


88  DUDLEY    MEMORIAL   VOLUME 

Sheaths  deciduous;  leaves  with  1  -vascular  bundle,  in  5s  (except  No.  6,  7) ; 

wood  light-colored  and  soft. 
Cones   cylindric;   scales  thin;   wings  elongated;    leaves  in   5s,   serrulate. 

White  Pines. 
Leaves  not  sharp-pointed;  resin  ducts  not  surrounded  by  strengthening 

cells. 

Leaves  grayish  green,  soft,  recurved  or  drooping;  branchlets  glabrous. 
Leaves  5-8  inches  long,  drooping;  cones  6-10  inches  long;  scales 

abruptly  pointed  at  apex.  1.  P.  excelsa. 

Leaves  2-4   (rarely  5)   inches  long;  cones  2-4  inches  long;  scales 

rounded  at  apex.  2.  P.  Strobus. 

Leaves  bluish   green,   stiff  and  erect;   branchlets  puberulent;   cones 

4-11  inches  long;  scales  pointed.  3.  P.  monticola. 

Leaves  sharp-pointed  and  stiff,  2-4  inches  long,  dark  bluish  green;  resin 
ducts  surrounded  by  strengthening  cells;  cones  12-20  inches  long. 

4.  P.  Lambertiana. 
Cones  not  cylindric;  scales  thickened;  wings  reduced  to  a  ring;  leaves  not 

serrulate.     Stone  Pines. 

Leaves  in  5s;  cones  2l/2-3  inches  long.  5.  P.  Cembra. 

Leaves  in  2s  or  4s;  cones  £4-2  inches  long. 

Leaves  in  4s.  6.  P.  quadrifolia. 

Leaves  in  2s.  7.  P.  edulis. 

Sheaths  persistent;  leaves  with  2  vascular  bundles,  in  2s  or  3s  (except  Tor- 

reyana) ;    wood   resinous.     Pitch    Pines. 

Leaves  in  5s,  about  10  inches  long.  8.  P.  Torreyana. 

Leaves  in  2s  or  3s. 
Leaves  in  3s. 

Cones  with  the  umbo  ending  in  a  stout,  hooked  projection ;  leaves  8  or 

more  inches  long. 
Leaves  grayish  green,  drooping;  cones  chocolate-colored,  ovate. 

9.  P.  Sabiniana. 
Leaves  dull  green,  spreading;  cones  light  brown,  cylindric-ovate. 

10.  P.  Coulteri. 

Cones  with  rounded  or  flattened  apophysis ;  umbo  with  or  without  a 

slender  prickle. 

Umbo  with  a  prominent  prickle ;  apophysis  but  little  thickened. 
Branchlets  glaucous ;  leaves  bluish  green. 

11.  P.  Jeffreyi. 
Branchlets  not  glaucous;  leaves  bright  yellowish  green. 

12.  P.  ponder osa. 


GYMNOSPERMS   ON    STANFORD   GROUNDS ABRAMS  89 

Umbo  without  or  with  a  rudimentary  prickle. 

Leaves  slender,  drooping,  8-10  inches  long;  cones  cylindric,  4-8 
inches  long;  apophysis  low-pyramidal. 

13.  P.  canariensis. 
Leaves  not  drooping,  4-6  inches  long,  dark  green;  cones  ovate 

and  unsymmetrical ;  apophysis  on  the  outside  rounded. 

14.  P.  radiata. 
Leaves  in  2s. 

Cones  unsymmetrical;   scales  much  enlarged  on  the  outside;  armed 

with  prominent  prickles.  15.  P.  muricata. 

Cones  nearly  symmetrical;  prickles  none  or  inconspicuous. 

Leaves  slender  and  flexible,   2-4  inches  long;  apophysis  flattened, 

deep,  lustrous  brown.  16.  P.  halepensis. 

Leaves  stout  and  rigid. 

Cones  and  leaves  4  inches  long  or  more. 

Seeds  ^  inch  long,  with  a  very  short  wing;   apophysis  low, 

somewhat  6-sided.  17.  P.  Pinea. 

Seeds  scarcely  ^   inch  long,   much  shorter  than  the  wings; 
apophysis  pyramidal  and  sharply  keeled. 

18.  P.  Pinaster. 

Cones  and  leaves  1^2-3  inches  long. 
Trees  of  good  size. 

Leaves  flattened;  cone-scales  with  a  flattened  apex  projecting 

beyond  the  dorsal  umbo.  19.  P.  sylvestris 

Leaves  rounded  on  the  back,   grooved  beneath,   2-3   inches 
long;  apex  of  scale  not  projecting. 

20.  P.  nigra. 

Low  dwarf  shrub  with  dense  foliage  of  dull  green  leaves. 

21.  P.  montana. 

1.     Pinus  excelsa  Wall. 

Branchlets  greenish  brown,  glabrous,  glaucous;  leaves  very  slender, 
flaccid,  drooping,  grayish  green,  6-8  inches  long;  cones  on  stalks  1-2  inches 
long,  cylindric,  6-10  inches  long;  scales  sharp-pointed;  seeds  Y$  inch  long; 
wings  1  inch  long,  acute. 

A  native  of  the  Himalaya  Mountains,  where  it  attains  a  height  of  150 
feet.  One  tree  is  in  the  Arboretum  northeast  of  the  Cactus  Garden. 

2.     Pinus  Strobus  L.     White  Pine. 

Branchlets  glabrous,  green  or  greenish  brown;  leaves  soft  and  very 
flexible,  2-5  inches  long,  light  bluish  green;  cones  on  stalks  l/2-\  inch  long, 


90  DUDLEY    MEMORIAL   VOLUME 

cylindric,    2-4    inches    long;    scales   oblong-obovate,    flexible;    seed   reddish 
brown,  mottled  with  black,  %  mcn  l°ng;  wings  4  times  as  long,  acutish. 

A  native  of  eastern  America,  extending  from  Newfoundland  to  Mani- 
toba, south  to  Georgia  and  Iowa.  Specimens  are  along  Pine  Avenue,  be- 
tween University  Avenue  and  the  automobile  road  and  in  the  nursery  south 
of  the  lath-house. 

3.     Pinus  monticola   Don.     Mountain  White   Pine. 

Branchlets  puberulent,  yellowish  or  reddish  brown;  leaves  stiff,  bluish 
green  and  glaucous,  1^-4  inches  long;  cones  short-stalked,  cylindric,  5-11 
inches  long,  yellowish  brown ;  scales  pointed  by  the  slightly  thickened  umbo ; 
seeds  ^  inch  long,  reddish  brown,  mottled  with  black;  wings  3  times  as 
long,  acute. 

A  native  of  western  America,  extending  from  British  Columbia  to  Idaho 
and  the  high  mountains  of  California.  One  tree  east  of  Cactus  Garden, 
a  few  others  are  scattered  through  the  Arboretum. 

4.     Pinus  Lambertiana  Dougl.     Sugar  Pine. 

Branchlets  pubescent,  brown;  leaves  ^  inch  long,  stout,  sharp-pointed, 
dark  bluish  green  with  conspicuous  white  lines  on  back;  cones  on  stalks, 
2-3l/2  inches  long,  cylindric,  light  brown,  shiny,  10-20  inches  long;  seed 
l/2  inch  long,  dark  brown  or  nearly  black;  wing  rounded  at  apex. 

A  Pacific  Coast  species,  extending  from  southern  Oregon  to  northern 
Lower  California.  The  largest  and  most  magnificent  of  all  the  pines. 
Several  trees  are  north  of  the  Angel  of  Grief,  and  one  or  two  fairly  large 
trees  are  north  of  the  Stanford  Residence. 

5.     Pinus  Cembra  L.     Swiss  Stone   Pine. 

Branchlets  with  yellowish  brown  tomentum;  leaves  straight,  dark  green 
on  back,  bluish  white  inside,  2-3^  inches  long;  cones  short-peduncled, 
ovate,  light  brown,  2l/2-Zl/2  inches  long;  scales  rounded  at  apex;  apophysis 
much  broader  than  high;  seed  l/2  inch  long. 

Native  of  the  Alps  and  extending  northward  to  Russia  and  northern 
Asia,  where  it  attains  70  or  occasionally  120  feet.  One  small  tree  which  has 
not  fruited  is  at  the  entrance  of  the  Cactus  Garden. 

6.     Pinus  quadrifolia  Parl.     Parry's  Pinon. 

Branchlets  puberulous,  light  grayish  brown;  leaves  3-5  (usually  4),  rigid, 
incurved,  light  green  on  back,  whitish  inside,  1^-2  inches  long;  cones  sub- 
globose,  \y2-1  inches  long,  chestnut  brown;  apophysis  thick,  pyramidal,  con- 
spicuously keeled:  umbo  with  minute  recurved  prickle;  seed  l/2  inch  long. 


GYMNOSPERMS   ON    STANFORD  GROUNDS ABRAMS  91 

Tree  attaining  40  feet,  with  spreading  branches  forming  a  rounded  top ; 
bark  dark  brown  tinged  with  red,  shallowly  fissured.  Native  of  the  extreme 
southern  part  of  California  and  extending  into  the  mountains  of  northern 
Lower  California.  There  are  no  specimens  of  this  tree  in  the  Arboretum, 
but  one  tree  which  fruited  in  1911  is  at  17  Salvatierra  Street,  and  another 
small  specimen  is  at  8  Alvarado  Row. 

7.     Pinus   edulis   Engelm.     Pinon. 

Leaves  in  2s  or  rarely  3s,  stout,  rigid,  incurved,  dark  green  on  the  back, 
marked  within  by  several  rows  of  stomata,  fa\l/2  inches  long,  persistent  for 
3  or  4  years  or  sometimes  longer;  staminate  flowers  dark  red;  cones  fall/2 
inches  long  and  nearly  as  broad;  seeds  ovate,  dark  red-brown  below,  orange- 
yellow  above,  %  inch  long ;  wings  ^  inch  wide. 

A  small  tree  with  a  divided  trunk,  30  to  40  feet  high.  Native  of  the 
southern  Rocky  Mountains,  extending  from  eastern  Utah  and  southwestern 
Wyoming  southward  to  the  mountains  of  northern  Mexico.  One  young  tree 
is  in  Professor  Durand's  garden. 

8.  Pinus  Torreyana  Parry.     Torrey  Pine. 

Branchlets  greenish  or  purplish,  glabrous;  leaves  rigid,  dark  green, 
8-12  inches  long;  cones  broadly  ovate,  4-6  inches  long,  chocolate-brown; 
apophysis  low-pyramidal;  umbo  elongated,  reflexed,  with  a  short  spiny  tip; 
seed  24  inch  l°ng;  short- winged. 

Tree  40  or  occasionally  60  feet  high,  with  spreading  branches  and  dark 
brown  bark.  Perhaps  the  rarest  pine,  known  only  in  two  small  groves: 
one  is  at  Del  Mar,  San  Diego  County,  the  other  is  on  Santa  Rosa  Island 
off  the  coast  of  southern  California.  Good-sized  trees  are  near  the  middle 
of  the  Arboretum  just  west  of  University  Avenue. 

9.  Pinus  Sabiniana  Dougl.     Digger  Pine. 

Leaves  slender,  drooping,  grayish  green,  8-12  inches  long;  cones  pendent 
on  stalks  2  inches  long,  light  red-brown,  6-10  inches  long;  apophysis 
pyramidal,  sharply  keeled,  flattened  at  the  straight  or  incurved  apex;  seeds 
24  inch  long;  short-winged. 

Tree  50  to  80  feet  high  with  the  trunk  usually  divided  into  several  stems, 
forming  a  round-topped  head.  Native  of  the  inner  Coast  Ranges  and  the 
foothills  of  the  Sierra  Nevada,  California.  Specimens  are  in  the  Roble  Gar- 
den, near  the  Museum,  and  in  the  Nursery. 

10.     Pinus   Coulteri   Don.     Coulter   Pine. 

Leaves  stout,  acuminate,  dark  bluish  green,  not  drooping,  6-12  inches 
long;  cones  short-stalked,  cylindric-ovate,  yellowish  brown,  9-14  inches  long; 


92  DUDLEY    MEMORIAL   VOLUME 

apophysis  elongated-pyramidal,  gradually  narrowed  into  straight  or  incurved 
umbo;  seed  y2  inch  long. 

Tree  80  feet  high,  forming  a  loose  pyramidal  head.  Native  of  the 
Coast  Ranges  of  California.  Trees  are  planted  in  the  Roble  Garden  near 
the  Museum  and  elsewhere. 

11.     Pinus  Jeffreyi  Murry.     Jeffrey's  Pine. 

Leaves  stout,  5-8  inches  long,  pale  bluish  green;  cones  conic-ovate, 
light  brown,  6-12  inches  long;  apophysis  depressed,  keeled;  umbo  elongated 
into  a  slender  recurved  spine;  seed  about  ^  inch  long. 

Forest  tree  attaining  a  height  of  nearly  200  feet,  with  short  spreading 
branches  forming  an  open  pyramidal  head.  Native  of  the  mountains  of 
Oregon  and  California.  Specimens  are  in  the  Nursery  south  of  the  lath- 
house. 

12.  Pinus  ponderosa  Dougl.     Western  Yellow  Pine. 
Branchlets   reddish   brown;    leaves   dark  yellowish   green,    5-10   inches 

long;  cones  ovate-oblong,  light  reddish  brown,  3^-5  inches  long;  apophysis 
flattened,  keeled;  umbo  triangular,  ending  in  a  stout,  straight  or  incurved 
prickle;  seed  %  inch  long;  wing  1  inch  long. 

Tree  150  or  occasionally  230  feet,  forming  a  spire-like  head.  Native 
of  western  America,  extending  from  British  Columbia  to  Mexico,  and  from 
western  Nebraska  and  Texas  to  California.  A  row  of  trees  is  west  of  the 
Nursery  lath-house,  others  are  between  University  Avenue  and  the  Mausoleum, 
and  a  few  large  trees  are  north  of  the  Stanford  Residence. 

13.  Pinus  canariensis  C.  Sm.     Canary  Island  Pine. 
Branchlets   yellowish;  leaves  slender,  8-11  inches  long,  light  green,  flexi- 
ble, usually  drooping,  flattened;  cones  ovoid,  4-7  inches  long,  light  brown, 
lustrous;    apophysis   low-pyramidal,   umbo   obtuse   or   sometimes   depressed; 
seed  ^2  inch  long. 

Tree  80  feet  high  with  slender  branches  forming  a  round-topped  head. 
Native  of  the  Canary  Islands  and  Teneriffe.  Several  trees  are  west  of  the 
Nursery  and  one  near  Mausoleum  Avenue.  Some  of  the  trees  west  of  the 
nursery  which  weite  severely  burned  three  or  four  years  ago  are  sending  out 
new  shoots  from  the  burned  branches  and  trunks.  The  primary  leaves  on 
these  shoots  are  silvery  glaucous. 

14.     Pinus  radiata  Gord.     Monterey  Pine. 

Leaves  bright  green,  3-6  inches  long ;  cones  light  brown,  lustrous,  conic- 
ovate,  unsymmetrical,  3-5  inches  long,  2-4  inches  broad;  scales  on  upper  sur- 


GYMNOSPERMS   ON    STANFORD   GROUNDS ABRAMS  93 

face  with  rounded  apophysis,  those  on  the  lower  surface  with  nearly  flat 
apophysis ;  prickles  very  minute ;  seeds  black,  %  m°h  l°ng>  half  the  length  of 
the  wing. 

Tree  80  to  100  feet,  broadly  pyramidal  or  with  round-topped  head.  Na- 
tive of  the  coast  of  California  and  abundant  at  Monterey.  Commonly  culti- 
vated on  the  University  grounds. 

15.  Pinus  muricata  Don.     Prickle-cone  Pine. 

Leaves  bright  green,  rather  stout,  4-6  inches  long,  strongly  serrate; 
cones  1-iy2  inches  long,  very  unsymmetrical  by  the  enlargement  of  scales  on 
outside;  prickles  prominent;  seeds  Ys  inch  long;  wings  y2  inch  long. 

A  compact  pyramidal  or  branching  tree  50  to  90  feet  high,  with  dense 
foliage.  Native  of  the  California  coast.  Specimens  are  among  the  pines 
between  the  Post  Office  and  the  Quadrangle.  Others  were  formerly  back  of 
the  Chemistry  Building. 

16.  Pinus  halepensis  Ait.     Aleppo  Pine. 

Leaves  dull  green,  very  slender  for  the  group,  2^2-3  inches  long,  tufted 
at  the  end  of  the  branches;  cones  conical,  smooth,  2^-3  inches  long,  deep 
lustrous  brown ;  scales  flattened,  ^4  inch  broad ;  apophysis  but  slightly  thick- 
ened, flat  and  smooth,  except  for  a  rather  faint  transverse  ridge. 

A  low  spreading  tree  with  an  open,  thin  and  straggling  aspect,  20-40  feet 
high.  Native  of  the  Mediterranean  region,  extending  from  Portugal  to  Asia 
Minor.  Several  trees  are  planted  in  the  Arboretum;  one  especially  fine 
specimen  is  between  the  Mausoleum  and  the  Museum. 

I6a.  Pinus  halepensis  Pityusa  Stevens. 

Larger  tree  than  the  typical  form,  with  leaves  4-5  inches  long  and  cones 
3-5  inches  long. 

Native  of  the  eastern  Mediterranean  region.  Specimens  are  south  of 
the  Nursery. 

17.  Pinus  Pinea  L.     Italian  Stone  Pine. 

Leaves  stout,  straight,  deep  shiny  green,  5-8  inches  long;  cones  5-6 
inches  long,  broadly  ovate,  lustrous  brown;  scales  thick  and  heavy,  1  inch 
broad;  apophysis  but  slightly  elevated,  often  6-sided;  umbo  with  a  short 
blunt  prickle;  seeds  fy  mcn  l°ngi  with  broad  but  very  short  wings. 

Tree  25-60  feet,  with  a  broad,  rounded  head.  Common  on  the  sandy 
shore  of  Tuscany.  One  tree  is  north  of  the  Nursery  lath-house. 

18.  Pinus  Pinaster  Ait.     Cluster  Pine. 

Leaves  glossy  green,  very  stout,  stiff  and  twisted,  acute,  5-9  inches  long, 
appearing  in  definite  and  somewhat  remote  bands;  cones,  in  the  older  trees, 


94  DUDLEY    MEMORIAL   VOLUME 

clustered,  conic-oblong,  4-7  inches  long;  apophysis  nearly  ^4  incn  broad,  low- 
pyramidal,  conspicuously  keeled  with  a  prominent  triangular  umbo;  seed  l/z 
inch  long ;  wing  about  1  inch  long. 

Tree  100  feet  high  with  regular  pyramidal  habit  and  rapid  growth. 
Native  of  the  coastal  region  of  southern  Europe.  A  variety  has  been  exten- 
sively used  in  Europe  to  reclaim  wastes  of  maritime  sands.  Several  trees  are 
in  the  row  of  pines  between  the  Post  Office  and  the  Quadrangle.  A  hand- 
some specimen  is  in  the  rear  of  Madrone  Hall,  and  a  few  trees  are  south  of 
the  Nursery. 

19.     Pinus  nigra  Arnold.     Austrian  Pine. 

Leaves  stout  and  very  rigid,  \l/2-1l/2  inches  long,  abruptly  sharp-pointed, 
rounded  on  the  back,  deeply  grooved  beneath,  dark  green;  cones  2^  inches 
long,  light  brown;  scales  about  1  inch  long,  ^  inch  broad;  apophysis  low- 
pyramidal,  slightly  keeled  and  with  a  minute  prickle. 

This  species  is  more  generally  known  under  the  name  Pinus  Austriaca. 
It  is  a  native  of  southeastern  Europe.  A  small  tree  is  southwest  of  the 
Nursery  lath-house. 

20.  Pinus  sylvestris  L.     Scotch  Pine. 

Leaves  light  green,  flattened  and  twisted,  stiff.  \-\l/2  inches  long;  cones 
2-2y2  inches  long,  light  brown;  scales  ^  inch  broad;  apophysis,  at  least  of 
the  middle  scales,  with  a  dorsal  knob-like,  keeled  and  truncated  thickening, 
and  a  thin,  somewhat  recurved  apex;  seed  l/\.  inch  long;  wing  narrow,  acute. 

A  forest  tree  70-100  feet  high,  with  pyramidal  head  and  scattered  foli- 
age. The  Scotch  Pine  is  one  of  the  important  timber  trees  of  Europe. 
Several  trees  are  in  the  Nursery  north  of  the  lath-house,  and  one  is  in  about 
the  middle  of  the  Arboretum  near  University  Avenue. 

21.  Pinus  montana  Mill.     Swiss  Mountain  Pine. 

Leaves  stout  and  crowded,  ^-2  inches  long,  bright  green,  acutish ;  cones 
^-2^4  inches  long,  light  grayish  brown;  apophysis  low-pyramidal,  with  a 
black  band  surrounding  the  umbo. 

A  variable  species,  often  a  low  dwarf  shrub,  or  sometimes  a  pyramidal 
tree  40  feet  high.  Native  of  the  mountains  of  central  and  southwestern 
Europe.  One  dwarf  specimen  is  on  the  west  side  of  the  Cactus  Garden. 

2.  Cedrus.     Cedars. 

Large  forest  trees,  with  stiff  4-sided,  more  or  less  sharply-pointed  leaves 
in  clusters  at  the  ends  of  short,  stubby  lateral  branchlets.  Flowers  monoecious, 
erect;  cones  ovate,  3-5  inches  long,  with  broad,  thin,  closely  imbricated 
scales. 


GYMNOSPERMS   ON    STANFORD   GROUNDS ABRAMS  95 

A   genus  of  three  closely  allied  species,   confined  to  northern  Africa, 
Asia    Minor   and   the    Himalaya    Mountains.     These    are   the   true   cedars, 
although  the  name  is  applied  to  a  number  of  totally  different  genera,  such  as 
Libocedrus,  Juniperus,  Thuja  and  Chamcecyparis. 
Branches  stiff  horizontal  or  ascending,  not  pendulous;   cones  truncate  and 

often  concave  at  apex. 
Branches  mostly  ascending;   leading  shoots  not  nodding;   leaves  mostly 

less  than  1  inch  long,  thicker  than  broad.          1.  C.  atlantica. 
Branches  horizontal;   leading  shoots  nodding;   leaves  1-1 J4  inches  long, 

broader  than  thick.  2.  C,  Libani. 

Branches  and  leading  shoots  pendulous;  cones  obtuse;  leaves  often  2  inches 
long.  3.  C.  Deodar  a. 

1.  Cedrus  atlantica  Manetti.     Mt.  Atlas  Cedar. 

Leaves  mostly  less  than  1  inch  long,  rigid,  dark  or  glaucous  green,  thicker 
than  broad ;  cones  2-3  inches  long,  light  brown. 

A  pyramidal  tree  120  feet  high,  with  ascending  branches  and  erect,  or  at 
least  not  drooping,  leading  shoots.  A  horticultural  variety,  glauca,  has 
leaves  very  glaucous  with  a  silvery  hue.  Native  of  the  mountains  of  northern 
Africa.  Frequently  planted  on  the  University  grounds.  A  handsome  speci- 
men of  the  variety  is  east  of  the  Cactus  Garden. 

2.  Cedrus  Libani  Barr.     Cedar  of  Lebanon. 

Leaves  1  inch  long  or  more,  dark  green  or  in  some  forms  bluish  green 
or  even  silvery,  broader  than  thick;  cones  3-4  inches  long,  brown. 

Large  forest  tree,  with  wide-spreading,  horizontal  branches  and  nodding 
leading  shoots.  Native  of  the  mountains  of  Syria  and  Asia  Minor.  Two 
trees  are  east  of  the  Cactus  Garden,  near  the  silver-leaved  Mt.  Atlas  Cedar, 
and  another  is  in  the  western  part  of  the  Nursery,  surrounded  by  a  cluster 
of  the  Mt.  Atlas  Cedar. 

3.  Cedrus  Deodara  Loud.     Deodar. 

Leaves  1-2  inches  long,  dark  bluish  green,  rigid,  as  thick  as  broad; 
cones  3^2-S  inches  long,  reddish  brown. 

Large  forest  tree,  often  150  feet  high,  with  branches  and  leading  shoots 
pendulous.  Several  horticultural  varieties  are  in  cultivation:  one,  argentea, 
has  silvery  leaves.  Native  of  the  Himalaya  Mountains.  Specimens  are  in 
Encina  and  Roble  Gardens,  and  the  variety,  argentea,  is  in  the  Arboretum 
north  of  the  Mausoleum. 


96  DUDLEY    MEMORIAL   VOLUME 

3.  Picea.     Spruce. 

Trees  with  spreading  or  sometimes  pendulous  branches.     Leaves  linear, 
flat  or  4-sided,  spirally  arranged,  sometimes  appearing  2 -ranked,   not  nar- 
rowed into  a  leaf -stalk ;  leaf -scars  raised  on  prominent  woody  pedicels,  which 
give  the  twigs  a  decided  roughness  after  the  leaves  have  fallen.     Cones  pen- 
dulous, their  scales  persistent,  completely  concealing  the  very  short  bracts. 
The  eighteen  known  species  are  confined  in  their  distribution  to  the  north 
temperate  and  subarctic  regions. 
Leaves  quadrangular  with  stomata  on  all  4  sides. 
Cone-scales  rounded  at  apex. 

Leaves  Y$~Y$  inch  long,  obtuse;  branchlets  not  long-pendulous. 
Young  twigs  glabrous;  cones  1^-2  inches  long. 

1.  P.  canadensis. 
Young  twigs  pubescent;  cones  3-5  inches  long. 

2.  P.  orientalis. 
Leaves  1^4-2  inches  long,  sharp-pointed.              3.  P.  Smithiana. 

Cone-scales  not  rounded  at  apex;  leaves  acute  or  sharp-pointed. 

Leaves   dark  green,   abruptly  acute;   cone-scales  firm  on  the  margins, 

truncate  at  apex.  4.  P.  excelsa. 

Leaves  blue  green,  acuminate  and  callous-tipped ;  cone-scales  rhomboidal, 

their  margins  flexuose.  5.  P.  Parry  ana. 

Leaves  flattened,  with  2  silvery  bands  of  stomata  above. 

6.  P.  sitchensis. 

1.  Picea  canadensis  (Mill.)  B.  S.  P.     White  Spruce. 

Leaves  spreading  from  all  sides  of  glabrous  twigs,  Y*~Y$  incn  long,  1/24 
inch  wide,  sharp-pointed,  dull  glaucous  green;  flowers  pale  red  or  yellowish; 
cones  1J4-2  inches  long,  about  1  inch  broad;  scales  rounded  at  apex,  Y$  inch 
wide. 

A  tree  50-150  feet  high,  with  a  symmetrical  pyramidal  head.  Native  of 
north  temperate  and  subarctic  America,  extending  from  Alaska  to  Labrador 
and  from  Montana  to  New  England.  One  tree  is  near  the  center  of  the 
Cactus  Garden. 

2.  Picea  orientalis  (L.)  Carr.     Caucasian  Spruce. 

Leaves  spreading  from  all  sides  of  the  pubescent  twigs,  Y$~Y*  incn 
long,  1/12  inch  wide,  4-sided,  blunt  at  apex,  dark  lustrous  green;  flowers 
carmine;  cones  3-5  inches  long,  about  \Yz  inches  broad;  scales  rounded  at 
apex,  Y$  inch  wide. 

Native  of  the  Caucasus  and  Asia  Minor.  Two  trees  are  near  the  center 
of  the  Cactus  Garden. 


GYMNOSPERMS   ON    STANFORD   GROUNDS ABRAMS  97 

3.  Picea  Smithiana  (Wall)   Bois.     Himalaya  Spruce. 

Leaves  spreading  from  all  sides  of  the  long  pendulous  branches,  \Y^-1 
inches  long,  1/20  inch  wide,  sharp-pointed,  light  green;  cones  5-7  inches 
long,  about  2  inches  broad;  scales  firm  and  shiny,  about  ^4  inch  wide, 
crowded  into  close  spirals,  rounded  at  apex. 

A  common  tree  in  the  coniferous  forests  of  the  Himalaya  Mountains. 
One  tree  is  on  the  west  side  of  the  Cactus  Garden  and  two  others  are  near 
the  Angel  of  Grief. 

4.  Picea  excelsa  Link.     Norway  Spruce. 

Leaves  spreading  from  all  sides  of  the  sparsely  pubescent  light  brown 
twigs,  1/2-^3  inches  long,  1/24  inch  wide,  4-sided,  sharp-pointed,  bright  green; 
cones  2^-5  inches  long,  about  half  as  broad;  scales  ^3  inch  wide,  their 
exposed  tips  3-sided,  the  2  lateral  margins  oblique  from  a  truncate  and  erose 
apex. 

Native  of  the  mountains  of  central  and  northern  Europe.  One  tree  is 
on  the  west  side  of  the  Cactus  Garden,  and  others  are  planted  elsewhere 
on  the  Campus. 

5.     Picea  Parryana  Sargent.     Blue  Spruce. 

Branchlets  glabrous;  leaves  stout,  rigid,  acuminate  and  sharp-pointed 
with  a  callous  tip,  fall/k  inches  long,  or  less  on  fertile  branches,  bluish 
green,  or  sometimes  silvery,  marked  on  both  sides  with  4-7  rows  of  stomata; 
staminate  flowers  yellow  tinged  with  red;  cones  oblong-cylindric,  about  3 
inches  long,  pale  chestnut  brown  and  glossy ;  scales  rhomboidal,  flexuose  on 
the  margins,  erose  at  the  summit;  seeds  y$  inch  long;  wings  twice  as  long. 

Forest  tree,  80  to  150  feet  high,  with  a  trunk  sometimes  3  feet  in 
diameter.  Native  of  the  mountains  of  Colorado,  eastern  Utah  and  Wyoming. 
A  young  tree  is  in  Professor  Durand's  garden. 

6.  Picea  sitchensis  Carr.     Tideland  Spruce. 

Leaves  spreading  from  all  sides  of  the  light  brownish  yellow  glabrous 
twigs,  green  and  rounded  below,  flat,  slightly  keeled  and  with  2  broad  silvery- 
bands  above,  sharply  acute,  y2-\  inch  long;  cones  cylindric-oval,  2^-4  inches 
long;  scales  rounded  and  erose  at  apex. 

Tree  100  to  200  feet  high,  with  slender  horizontal  branches.  Native  of 
the  northwest  coast,  extending  from  Alaska  to  northern  California.  One 
tree  is  in  the  Arboretum  between  the  the  Automobile  road  and  the  carline  near 
the  old  Sempervirens  avenue.  This  tree  was  apparently  identified  as  Picea 
Parryana  in  Dudley's  key. 


98  DUDLEY    MEMORIAL   VOLUME 

4.  Pseudotsuga. 

Large  evergreen  forest  trees,  with  flat,  indistinctly  2-ranked  leaves,  nar- 
rowed at  base  into  short  petiole;  leaf-scars  sessile.  Cones  pendent,  their 
3-pronged  bracts  much  exserted ;  seeds  prominently  winged. 

A  genus  of  three  species;  two  in  western  America,  and  one  in  Japan. 

1.  Pseudotsuga  taxifolia  Britton.     Douglas  Fir. 

Leaves  ^-\l/2  inches  long,  obtuse,  dark  green  above,  paler  beneath; 
cones  oblong-ovate,  2-4  inches  long,  light  reddish  brown,  with  broad,  rounded 
scales  and  well  exserted  3-parted  bracts. 

The  Douglas  Fir  is  the  most  important  timber  tree  of  western  America. 
It  often  attains  a  height  of  200  feet  or  even  more,  and  a  trunk  12  feet  in 
diameter.  Frequently  planted  in  the  Arboretum.  Native  trees  are  growing 
on  the  Palo  Alto  estate  at  the  north  end  of  Jasper  Ridge. 

5.  Abies.     Firs. 

Trees  generally  pyramidal  in  outline,  with  their  branches  in  whorls. 
Leaves  solitary,  scattered  along  the  twigs  in  definite  spirals,  but  often  appear- 
ing 2-ranked  by  a  twist  near  the  base,  linear,  flat  or  slightly  4-sided,  some- 
times narrowed  at  base  but  not  into  a  distinct  leaf-stalk;  leaf -scars  sessile  on 
the  smooth  twigs.  Cones  erect  on  the  branches,  their  scales  deciduous,  and 
bracts  exserted  beyond  scales  or  concealed  by  them. 

A  genus  of  24  known  species,  which  inhabit  the  northern  or  mountainous 
parts  of  the  northern  hemisphere. 
Leaves  stomatiferous  and  alike  in  color  on  both  surfaces. 

Leaves  rigid,  spreading  from  all  sides,  Ys'Y*  inch  long,  closely  crowded  on 
the  twigs,  their  much  enlarged  bases  nearly  contiguous. 

Leaves  Ys'Y*  inch  long,  steel  green,  1/12  inch  wide,  somewhat  4-angled. 

1.  A.  pinsApo. 
Leaves  Y*~2A  mcn  l°ng>  dark  green,  Y%  inch  wide. 

2.  A.  barborensis. 
Leaves   curved  upward  on  twigs,   more  or  less   2-ranked,   pale  glaucous 

green,  1-2  inches  long.  3.  A.  concolor. 

Leaves  stomatiferous  only  on  lower  surface. 

Leaves  bright  green  on  lower  surface,  distinctly  2-ranked. 

4.  A.  Momi. 

Leaves  silvery  on  lower  surface  with  2  broad  bands  of  stomata. 
Leaves  not  pungently  acute. 
Leaves  obtuse. 

Leaves  distinctly   2-ranked,   forming  flat  sprays;   twigs  ill-scented 
when  bruised.  -5.  A.  grandis. 


GYMNOSPERMS   ON    STANFORD   GROUNDS ABRAMS  99 

Leaves  mostly  curved  upward;  twigs  not  ill-scented. 

Cones  6  inches  long  or  less;   bracts  projecting  beyond  scales; 
leaves  ^j-1  inch  long,  bright  green  on  upper  surface. 

6.  A.  nordmanniana. 
Cones  8  inches  long;  bracts  concealed  by  scales;   leaves   1-1  j4 

inches  long,  dark  green  on  upper  surface. 

7.  A.  cilicia. 
Leaves  acute.                                                        8.  A.  cephalonica. 

Leaves  pungently  acute;  bracts  long-exserted.         9.  A.  venusta. 

1.     Abies  pinsapo  Boiss.     Spanish  Fir. 

Leaves  dark  steely  green  on  both  surfaces,  rigidly  spreading  from  all 
sides  of  the  twig,  crowded,  their  much  enlarged  bases  nearly  contiguous, 
y$-l/2  inch  long,  1/12  inch  wide,  slightly  keeled  on  both  surfaces,  stoma- 
tiferous  on  both  surfaces,  with  inconspicuous  gray  bands,  obtuse  but  curved 
upward  at  tip  and  appearing  acute;  cones  about  8  inches  long,  1^2-1% 
inches  broad;  scales  wedge-shaped,  1  inch  wide;  bracts  short,  rounded,  not 
stalked. 

The  Spanish  Fir  is  a  native  of  central  and  southern  Spain,  where  it 
forms  extensive  forests  on  the  mountains  at  altitudes  of  4,000-6,000  feet. 
Excellent  specimens  of  this  species  are  at  the  southeastern  border  of  the 
Cactus  Garden. 

2.  Abies  barborensis  M'Nab.     Algerian  Fir. 

Leaves  dark  lustrous  green  on  both  surfaces,  rigidly  spreading  from  all 
sides  of  twig,  enlarged  at  base,  less  crowded  than  in  the  preceding;  J4-/4 
inch  long,  ^  inch  wide,  rounded  at  apex,  flat  but  keeled  on  both  surfaces, 
stomatiferous  bands  inconspicuous  on  both  surfaces;  cones  10  inches  long, 
dull  grayish  brown;  scales  rounded  at  apex,  tapering  to  base,  1-1^  inches 
wide;  bracts  short,  rounded. 

Native  of  Mt.  Barbor  and  Mt.  Tababor,  Algeria,  where  it  is  found  with 
the  Mt.  Atlas  cedar.  It  is  a  forest  tree,  often  60  feet  high,  with  compact 
branches  forming  symmetrical  pyramidal  heads.  Two  trees  are  along  the 
eastern  border  of  the  Cactus  Garden  near  Abies  grandis. 

3.  Abies  concolor  (Gord.)  Parry.     White  Fir. 

Leaves  ascending  or  indistinctly  2-ranked,  glaucous  green  and  stoma- 
tiferous on  both  surfaces,  1-2  inches  long,  1/12  inch  wide,  obtuse  or  acutish, 
faintly  grooved  above,  keeled  below,  with  2  lateral  shallow  grooves;  cones 
6-10  inches  long,  1-1^4  inches  broad;  scales  broader  than  long,  rounded  at 
apex,  gradually  tapering  to  the  base;  bracts  half  the  length  of  scales,  oblong, 


100  DUDLEY    MEMORIAL   VOLUME 

denticulate  and  mucronate  at  the  emarginate  or  truncate  apex;  seeds  3/12- 
4/12  inch  long,  dark  dull  brown;  wings  rose-colored,  widest  near  the  middle. 
A  large  forest  tree,  sometimes  200  feet  or  more  high,  with  a  trunk  6 
feet  in  diameter,  dull  grayish  brown  bark  and  short  stout  horizontal  branches 
in  remote  whorls.  Native  of  western  America,  extending  from  southern 
Oregon  to  the  high  mountains  of  northern  Lower  California  and  eastward 
to  New  Mexico.  A  number  of  young  trees  are  scattered  along  Mausoleum 
Avenue,  and  others  are  planted  elsewhere  on  the  Campus. 

4.  Abies  Momi  Siebold.     Momis  Fir. 

Leaves  appearing  2-ranked,  flat,  ^-1^4  inches  long,  1/12-1/8  inch  wide, 
grooved  and  bright  green  above,  faintly  keeled  and  bright  yellowish  green 
beneath,  inconspicuously  emarginate;  cones  2-3  inches  long;  scales  trans- 
versely oval  or  reniform,  \y2  inches  wide,  y$  inch  long,  lateral  margins  den- 
tate; bracts  stalked,  ^  as  long  as  scales;  spatulate,  mucronately  pointed. 

The  Momis  Fir  is  a  Japanese  species  and  is  often  cultivated  in  Japanese 
gardens  and  about  the  temples,  where  trees  over  100  feet  high  and  6  feet  in 
diameter  are  to  be  found.  One  small  tree  is  north  of  the  Cactus  Garden 
and  another  is  just  back  of  the  Angel  of  Grief. 

5.  Abies  grandis  (Dougl.)  Lindl.     Lowland  Fir. 

Leaves  appearing  2-ranked,  forming  a  flat  spray,  y$-\Yz  inches  long, 
thin  and  flexible,  deeply  grooved  and  dark  lustrous  green  above,  silvery  white 
below,  with  2  broad  bands  of  stomata,  emarginate;  cones  cylindric,  slightly 
narrowed  at  rounded  apex,  2-3^  inches  long,  1-1^  inches  broad;  scales 
broader  than  long,  rounded  at  apex,  abruptly  or  gradually  narrowed  to  the 
stalk-like  base;  bracts  about  %  the  length  of  scales,  obcordate,  irregularly 
serrate,  mucronate;  seeds  y2  inch  long,  light  brown;  wings  Y^A  inch  long. 

A  large  forest  tree,  sometimes  attaining  a  height  of  250  feet  and  a  diam- 
eter of  4  to  5  feet,  clothed  with  long  downward  sweeping  branches.  The 
wood  is  ill-scented  and  is  known  among  lumbermen  as  Stinking  Fir.  Native 
of  northwestern  America,  ranging  from  Vancouver  Island  southward  along 
the  lowlands  of  the  coast  to  Mendocino  County,  California,  and  eastward 
to  the  Bitterroot  Mountains  of  western  Montana  and  the  Blue  Mountains  of 
eastern  Oregon.  One  tree  is  situated  on  the  western  border  of  the  Cactus 
Garden  and  a  number  of  young  trees  are  planted  elsewhere  in  the  Arboretum. 

6.    Abies  Nordmanniana   (Stevens)   Spach.     Caucasian  Fir. 
Leaves  curved  upward,    24-1^4   inches  long,    1/12   inch  wide,   obtuse, 
slightly  grooved  and  dark  shiny  green  above,  keeled,  and  with  2  broad  sil- 
very bands  of  stomata  below;  cones  about  8  inches  long,  1^4  inches  broad; 


GYMNOSPERMS   ON    STANFORD   GROUNDS ABRAMS  101 

scales  1^2  inches  wide,  lateral  margins  lobed  near  the  apex,  serrate,  tapering 
to  a  short  slender  stalk;  bracts  1^4  inches  long,  the^v, recurved,  mucronate 
apex  exserted  beyond  scales.  JL-  &»  > 

The  Caucasian  or  Nordmann  Fir  is  a  native  of  Crfi|t|a  ai^the/fea,ucasus, 
where  it  inhabits  the  mountains  at  about  2,000  feet.     It 
sometimes  attaining  150  feet  in  height  and  6  feet  in  dii 
some  trees  are  in  the  Cactus  Garden,  and  other  smaller  trees  are' 


the  University  grounds.  ''/&/•    f/C/ 

7.  Abies  cilicia   (Kotsch.)    Carr.     Cilician  Fir. 

Leaves  curved  upward,  closely  resembling  those  of  the  preceding,  but 
darker  green  above  and  usually  longer,  1-1^4  inches;  cones  about  10  inches 
long,  1^4  inches  broad;  bracts  not  exserted. 

The  Cilician  Fir  with  the  Cedar  of  Lebanon  forms  extensive  forests  in 
the  Cilician  Taurus  at  elevations  of  4,000-6,000  feet.  Two  large  trees  are 
near  the  center  of  the  Cactus  Garden. 

8.  Abies  cephalonica  Loud.     Cephalonian  Fir. 

Leaves  spreading  at  right  angles  from  all  sides  of  the  twig,  dark  shiny 
green  above,  with  2  silvery  bands  beneath,  tapering  from  base  to  the  sharp- 
pointed  apex,  their  bases  dilated  lengthwise ;  cones  cylindric,  5-6  inches  long ; 
scales  rounded  above  and  entire,  wedge-shaped  toward  base;  bracts  projecting 
beyond  scales,  linear-oblong,  unequally  toothed  at  apex. 

Native  of  the  island  of  Cephalonia,  Greece.  One  specimen  is  on  the 
southwest  border  of  the  Cactus  Garden. 

9.  Abies  venusta  (Dougl.)   Koch.     Santa  Lucia  Fir. 

Leaves  appearing  2-ranked,  1-1^4  inches  long,  ^  inch  wide,  tapering  at 
both  ends  and  ending  in  a  sharp  spiny  tip  at  apex,  bright  green  above,  gray- 
ish beneath  with  2  broad  bands  of  stomata;  cones  ovate,  3-4  inches  long; 
bracts  much  exceeding  the  scales. 

Native  of  the  Santa  Lucia  Mountains.  Tree  often  100  to  150  feet  high, 
narrowed  toward  the  top  into  a  spire-like  head.  A  young  tree  is  on  the  left 
hand  side  of  the  Palo  Alto  entrance  to  the  Campus,  another  is  in  Encina 
Garden. 

Tribe  3.  Taxodeae. 

Leaves  and  floral  parts  spirally  arranged,  or  the  leaves  whorled  in 
S  dado  pit ys ;  ovuliferous  scales  bractless,  forming  woody  cones;  ovules  usu- 
ally several  to  each  scale  and  erect;  seeds  small,  sharply  and  irregularly 
angled. 


102  DUDLEY    MEMORIAL   VOLUME 

Leaves  in  whorls,  elongated.  6.  Sciadopitys. 

Leaves  spirally  arranged  or  appearing  2-ranked. 

Leaves  persistent  for  more  than  one  season ;  seeds  several  to  a  scale. 
Cone-scales  with  several  sharp  projections.        7.  Sequoia. 
Cone-scales  without  sharp  points.  8.  Cryptomeria. 

Leaves  deciduous  with  the  slender  twigs,  or  sometimes  persistent  for  a  year ; 
seeds  2  to  a  scale.  9.  Taxodium. 

6.    Sciadopitys. 

Evergreen  pyramidal  tree,  with  long  needle-like  leaves  in  whorls.  Flow- 
ers monoecious;  staminate  with  spirally  arranged  2-celled  anthers.  Cones 
ovate-oblong,  with  broadly  orbicular  scales. 

1.  Sciadopitys  verticillata  Sieb.   &  Zucc.     Umbrella   Pine. 

Leaves  in  whorls  of  15-30,  3-6  inches  long,  grooved  on  both  surfaces, 
dark  green  above,  and  with  white  band  beneath;  cones  3-4  inches  long. 

This  species,  which  is  the  only  member  of  the  genus,  is  a  native  of 
Japan.  One  young  specimen  is  temporarily  placed  in  the  Nursery  lath- 
house. 

7.  Sequoia. 

Tall  massive  forest  trees  with  trunks  usually  heavily  buttressed  at  base, 
covered  with  thick  fibrous  bark.  Leaves  evergreen,  linear  or  scale-like,  de- 
current  on  the  twigs.  Flowers  terminating  the  branchlets,  monoecious ;  pollen- 
sacs  several  on  the  lower  half  of  the  connective.  Cones  maturing  the  first 
year,  with  spirally  arranged,  peltate  scales;  seeds  several  to  each  scale,  red- 
dish brown. 

Once  a  genus  of  several  species  widely  distributed  over  North  America, 
Europe  and  Asia,  but  now  reduced  to  two  and  restricted  to  the  Coast  Ranges 
and  the  Sierra  Nevada  of  California. 
Leaves  of  2  kinds,  ordinarily  flat,  linear  and  2-ranked,  but  on  leading  shoots 

often  scale-like  and  spreading  from  all  sides  of  the  branchlets. 

1.  S.  sempermrens. 
Leaves  all  scale-like  and  scattered  on  all  sides  of  the  branchlets. 

2.  S.  Washingtoniana. 

1.  Sequoia  sempervirens  Endl.     Redwood. 

Leaves  distichously  spreading,  about  ^4  mcn  l°ng»  or  those  on  leading 
shoots  smaller  and  scale-like;  cone  oblong,  £4-1  inch  long. 

Tall,  magnificent  forest  trees  often  200-300  feet  high.  The  tallest  tree 
authentically  measured  is  340  feet.  Native  of  the  fog  belt  of  the  California 
Coast  Ranges,  extending  from  southwestern  Oregon  to  the  Santa  Lucia 
Mountains  in  Monterey  County,  and  inland  not  more  than  20  to  30  miles. 


GYMNOSPERMS   ON    STANFORD   GROUNDS ABRAMS  103 

Frequently  planted  on  the  University  grounds  but  enduring  badly  in  the  dry 
soil.  Native  trees  are  on  the  estate  along  San  Francisquito  Creek.  A  notable 
individual  is  the  "Palo  Alto"  at  the  railway  bridge  between  Palo  Alto  and 
Menlo  Park. 

2.    Sequoia  Washingtoniana  Sudw.     Giant  Sequoia. 

Leaves  scale-like,  arising  from  all  sides  of  the  branchlets,  y%-l/±  inch 
long,  glaucous  green;  cone  ovate-oblong,  2-3^2  inches  long. 

The  Giant  Sequoia  is  the  largest  and  probably  the  oldest  of  trees.  It 
is  200-325  feet  high  and  attains  a  diameter  of  30  feet.  Native  of  the  western 
slopes  of  the  Sierra  Nevada,  where  it  usually  occurs  in  small,  isolated  groves, 
but  it  forms  rather  extensive  forests  in  the  basins  of  the  Tulare,  Kings 
and  Kaweah  rivers.  Frequently  planted  on  the  University  grounds,  and 
apparently  better  adapted  to  our  local  climate  and  soil  than  the  Redwood. 

8.  Cryptomeria. 

Pyramidal  tree  with  reddish  brown  bark.  Leaves  linear-subulate,  aris- 
ing from  all  sides  of  the  twigs,  decurrent.  Flowers  small,  monoecious ;  stami- 
nate  oblong;  pistillate  globose.  Cones  globose,  with  thick,  wedge-shaped 
scales  furnished  with  a  recurved  projection  on  the  back  and  pointed  lobes  at 
the  apex. 

A  monotypic  Japanese  genus. 

1.  Cryptomeria  japonica  Don. 

Leaves  compressed,  somewhat  incurved,  y2-\  inch  long;  cones  reddish 
brown. 

A  handsome  evergreen  tree  extensively  planted  in  Japan,  especially  about 
temples.  A  few  trees  are  planted  on  the  University  grounds,  but  it  does  not 
thrive  in  our  dry  soil.  An  excellent  specimen  is  at  9  Lasuen  St. 

la.     Cryptomeria  japonica  elegans  Beissn. 

Low  dense  tree  or  shrub,  with  horizontal  branches  and  pendulous 
branchlets;  leaves  linear,  flattened,  soft,  bright  green  changing  to  bronze  in 
winter. 

A  garden  form.  Good  specimens  are  on  the  grounds  of  the  Stanford 
Residence. 

9.  Taxodium. 

Tall,  deciduous  or  evergreen  trees,  with  cinnamon  brown,  flaky  bark. 
Leaves  linear,  2-ranked,  falling  off  in  the  autumn  or  the  second  year  with  the 
short  slender  lateral  twigs ;  flowers  monoecious,  the  staminate  with  4-5  pollen- 
sacs  to  each  anther,  the  pistillate  solitary  or  in  pairs  at  the  ends  of  the  branch- 


104  DUDLEY  MEMORIAL  VOLUME 

lets  of  the  previous  year;  cones  globose  or  nearly  so,  maturing  the  first  year, 
their  scales  spirally  arranged,  thickened  at  apex  and  mucronate;  seeds  2  to 
each  scale,  triangular,  winged. 
Leaves  deciduous;  flowers  appearing  in  the  spring. 

1.  T.   distichum. 
Leaves  persistent;  flowers  appearing  in  the  autumn. 

2.  T.  mucronatum. 

I.  Taxodium  distichum  Rich.     Bald  Cypress. 

Leaves  deciduous  in  the  autumn,  narrowly  linear,  light  green,  Yz-'fy 
inch  long;  staminate  flowers  purplish,  in  panicles  4-5  inches  long;  cones  1 
inch  long,  without  mucros  at  maturity. 

Tall  tree  becoming  150  feet  high,  with  a  buttressed  trunk,  sometimes  12 
feet  in  diameter.  Native  of  the  southeastern  United  States.  A  handsome 
specimen  of  this  very  ornamental  tree  is  near  the  Angel  of  Grief,  and  another 
young  tree  is  at  23  Salvatierra. 

2.  Taxodium  mucronatum  Tenore.     Mexican  Bald   Cypress. 

Similar  to  the  last  but  leaves  persistent  through  the  winter;  flowers  ap- 
pearing in  the  autumn  instead  of  the  spring;  pollen-sacs  7-9  instead  of  4-5. 

Native  of  eastern  and  southern  Mexico.  Several  notably  large  trees  are 
known;  one  of  these,  the  Cypress  of  Montezuma,  was  a  noted  tree  four  cen- 
turies ago.  A  young  specimen  that  is  withstanding  our  winters  badly,  is  in 
Professor  Durand's  garden.  This  specimen  has  now  passed  through  three 
winters,  and  although  the  tips  of  the  young  shoots  have  been  killed  by  frost, 
the  leaves  have  retained  their  persistent  character. 

Tribe  4.  Cupresseae. 

Leaves  and  floral  parts  decussately  opposite  or  ternate,  the  former  scale- 
like   or  sometimes   subulate.     Ovuliferous  scales  woody   or   coalescent   and 
fleshy;  ovules  2-many  to  each  scale. 
Cones  woody. 

Scales  flat  or  wedge-shaped,  imbricate. 

Seeds  4-5  to  each  scale;  branchlets  frond-like;  leaves  with  broad  scales. 

10.  Thuyopsis. 
Seeds  usually  2  (1-3)  to  each  scale. 

Leaves  appearing  in  whorls  of  4,  flat.  11.  Libocedrus. 

Leaves  decussately  opposite.  12.  Thuya. 

Scales  peltate. 

Cones  maturing  the  second  year;  seeds  many  to  each  scale. 

13.  Cupressus. 


GYMNOSPERMS   ON    STANFORD  GROUNDS ABRAMS  105 

Cones  maturing  the  first  year ;  seeds  few  to  each  scale. 

14.  Chamcecy paris. 
Cones  becoming  fleshy  and  berry-like.  15.  Juniperus. 

10.  Thuyopsis. 

Evergreen  forest  tree  with  a  pyramidal  head.  Leaves  decussately  oppo- 
site, convex  above  and  somewhat  sac-like.  Flowers  monoecious,  solitary  and 
terminal;  staminate  cylindric;  cone  subglobose,  with  8-10  wedge-shaped 
scales;  seeds  5  to  each  scale,  compressed. 

A  monotypic  Japanese  genus. 

1.    Thuyopsis  dolabrata  Sieb.  &  Zucc.     Hatchet-leaved  Arborvitae. 

Leaves  glossy  green  above,  silvery  white  beneath,  rounded  at  apex,  the 
upper  and  lower  appressed,  the  lateral  spreading,  hatchet- shaped ;  cones  ^ 
inch  long ;  scales  reflexed  at  apex. 

A  native  of  canyons  and  moist  slopes  of  Japan.  Two  young  trees  are 
at  9  Lasuen  St. 

11.  Libocedrus. 

Evergreen  aromatic  trees,  with  fibrous  bark  and  flattened  spray-like 
branchlets.  Leaves  scale-like,  imbricate  in  4  rows.  Flowers  monoecious; 
staminate  with  12-16  stamens;  pollen-sacs  4.  Cones  oblong,  with  6  scales, 
the  lower  pair  much  reduced,  only  the  middle  pair  fertile;  seeds  2  to  each 
scale,  winged. 

A  genus  of  about  8  species ;  one  is  a  Calif ornian  species,  the  others  are 
in  western  South  America,  New  Zealand,  New  Caledonia,  New  Guinea,  For- 
mosa and  southwestern  China. 

1.  Libocedrus  decurrens  Torr.     Incense  Cedar. 

Leaves  yellowish  green,  appearing  in  whorls  of  4,  the  lateral  nearly  cov- 
ering the  obscurely  pitted  inner  ones;  cones  ^  incn  l°ng»  about  y$  inch 
thick;  scales  with  a  short  recurved  mucro. 

Tree  sometimes  150  feet  high,  with  an  irregularly  lobed  trunk  tapering 
from  a  broad  base,  and  a  reddish  brown,  fibrous  bark.  Native  of  the  Sierra 
Nevada  and  the  inner  Coast  Ranges,  extending  from  southern  Oregon  to 
northern  Lower  California.  The  row  of  trees  back  of  the  Engineering  Build- 
ings is  of  this  species.  Other  trees  are  planted  elsewhere  on  the  University 
grounds. 

12.  Thuya. 

Evergreen  trees  with  thin,  scaly  bark  and  decussate  scale-like  leaves. 
Flowers  monoecious;  staminate  ovoid,  with  4-6  pollen-sacs.  Cones  ovoid- 


106  DUDLEY    MEMORIAL   VOLUME 

oblong,  erect  or  drooping,  maturing  in  one  season;  scales  8-12,  the  middle 
2  or  3  pairs  fertile. 

Four  species  are  recognized  in  this  genus,  two  in  North  America  and  two 
in  Asia. 
Cones  pendulous ;  scales  thin,  with  a  minute  mucro. 

1.  T.  plicata. 
Cones  erect ;  scales  angled  on  the  back,  with  a  stout,  recurved  dorsal  hook. 

2.  T.  orientalis. 

1.  Thuya  plicata  Don.     Western  Red  Cedar. 

Leaves  bright  green  and  glossy  above,  dark  green  beneath,  with  whitish 
triangular  markings;  cones  cylindric-ovoid,  scarcely  over  l/2  inch  long; 
scales  8-10,  elliptic-oblong,  usually  the  3  middle  pairs  fertile;  seeds  com- 
pressed, notched  at  the  apex,  with  2  narrow  wings. 

A  large  forest  tree,  200  feet  high,  with  short  horizontal  branches  often 
with  pendulous  tips.  Native  of  northwestern  America,  extending  from 
Alaska  to  northern  California.  A  specimen  is  in  the  eastern  part  of  Encina 
Garden  and  another  is  near  Mausoleum  Avenue. 

2.  Thuya  orientalis  L.     Arborvitae. 

Leaves  acute,  bright  green,  with  a  small  dorsal  gland;  cones  erect, 
globose-ovate,  ^4  mcn  l°ngj  scales  usually  6,  oval;  angled  on  the  back,  and 
with  a  dorsal  horn-like  process,  the  uppermost  pair  sterile;  seeds  wingless. 

Pyramidal  trees  25  feet  high  or  bushy.  Native  of  Persia  and  extending 
to  eastern  Asia.  There  are  many  garden  forms  of  this  extensively  cultivated 
species.  Abundantly  planted  on  the  University  grounds. 

12.  Cupressus. 

Evergreen   trees    or    rarely    shrubs.     Leaves    decussately   opposite,    ap- 
pressed,  small,  scale-like.     Flowers  monoecious,  very  small,  terminating  short 
branchlets.     Cones  globose  or  nearly  so,  consisting  of  3-7  pairs  of  peltate 
woody  scales,  each  bearing  many  seeds,  maturing  the  second  year. 
Branchlets  terete  or  quadrangular,  not  forming  flat  spays. 
Leaves  glandless  or  with  an  obscure  dorsal  gland. 

Branchlets  erect  or  spreading,  the  ultimate  rather  short  and  stout. 
Ultimate  branchlets  terete  or  nearly  so ;  cones  usually  1  ^4  inch  long 

or  more.  1.  C.  sempervirens. 

Ultimate  branchlets  4-sided;  cones  globose,  1  inch  or  less  broad. 
Staminate  cones  subglobose,  with  2  lateral  anthers  in  each  row ; 
seeds  dull  grayish  or  blackish  brown. 

2.   C.  macro  car  pa. 


GYMNOSPERMS   ON    STANFORD  GROUNDS ABRAMS  107 

Staminate  cones  oblong-ovoid,  with  3  anthers  to  each  row;  seeds 

reddish  brown.  3.  C.  Goveniana. 

Branchlets  usually  pendulous,   the  ultimate  terete,   slender,   elongated; 

cones  globose,  l/2  inch  broad.  4.  C.  torulosa. 

Leaves  conspicuously  glandular  and  very  fragrant. 

5.  C.  Macnabiana. 
Branchlets  flattened,  forming  frond-like  sprays  on  long,  pendulous  branches. 

6.  C.  funebris. 

1.  Cupressus  sempervirens  L.     Italian  Cypress. 

Tree  attaining  80  feet,  the  typical  form  with  erect  branches  forming  a 
narrow  columnar  head;  leaves  closely  appressed,  ovate,  glandless  or  with 
inconspicuous  dark  pits;  staminate  flowers  cylindric,  with  4  lateral  anthers 
in  each  row;  cones  oblong  or  subglobose,  1*4  inches  long  or  more,  glossy; 
seeds  nearly  *4  mcn  l°ng>  reddish  brown,  with  light-colored  hilum. 

The  columnar  form  is  the  classical  Cypress  of  the  Greeks  and  Romans, 
and  is  much  cultivated  in  southern  Europe.  It  is  the  type  of  the  species, 
but  is  not  now  known  in  the  wild  state.  Commonly  planted  on  the  Univer- 
sity grounds. 

la.     Cupressus  sempervirens  horizontalis  Gord. 

Branches  horizontal,  forming  a  broad  pyramidal  head,  otherwise  like 
the  typical  form. 

This  form  is  also  common  on  the  Campus. 

2.  Cupressus  macrocarpa  Hartw.     Monterey  Cypress. 

Branchlets  short  and  stout,  arising  from  all  4  sides  of  the  twig;  leaves 
dark  green,  glandless  or  glands  marked  by  a  dark  pit;  rhombic-ovate,  acut- 
ish;  staminate  flowers  subglobose,  with  2  lateral  anthers  in  each  row;  cones 
globose  to  oblong,  ^4-lj^2  inches  long;  scales  8-12,  with  a  short  obtuse  umbo ; 
seeds  sharply  angled,  3/16  inch  long,  chestnut  brown,  with  a  conspicuous 
light-colored  hilum. 

Tree  attaining  40  feet  or  occasionally  70  feet,  broadly  oval  in  outline, 
with  spreading  branches  or,  especially  in  its  native  habitat,  with  horizontal 
branches  forming  a  broad  flat-topped  head.  The  Monterey  Cypress,  which 
is  the  most  abundantly  planted  tree  on  the  University  grounds,  is  restricted 
in  its  wild  state  to  two  small  groves  occupying  the  two  promontories,  Point 
Lobos  and  Cypress  Point,  that  mark  the  boundary  of  Carmel  Bay. 

3.     Cupressus  Goveniana  Gord. 

Branchlets  arising  from  all  4  sides  of  the  twigs,  quadrangular;  leaves 
ovate,  acute,  glandless  or  with  an  inconspicuous  dark  pit;  staminate  flowers 


108  DUDLEY    MEMORIAL   VOLUME 

oblong,  with  3  lateral  anthers;  cones  subglobose  or  oblong,  l/2-^  inch  broad; 
seeds  reddish  brown  with  hilum  of  similar  color. 

Tree  attaining  50  feet,  with  slender  spreading  or  erect  branches,  form- 
ing a  broad,  open,  or  pyramidal  head.  Native  of  the  coastal  region  of  north- 
ern and  central  California.  Two  trees  are  on  either  side  of  the  first  cross 
street  between  the  Library  and  the  Gymnasium. 

4.  Cupressus  torulosa  Don.     Himalayan  Cypress. 

Ultimate  branchlets  usually  arising  from  only  2  sides  of  the  twig,  terete, 
slender;  leaves  ovate,  acute,  bluish  green;  staminate  flowers  cylindric,  with 
3-4  lateral  anthers  in  each  row;  cones  globose,  y2  inch  broad;  scales  8-10 
with  a  small,  sharp  umbo;  seeds  light  brown,  glossy,  flattened  and  broadly 
winged. 

Tall,  slender  tree,  attaining  150  feet,  with  short  horizontal  branches  and 
slender  pendulous  branchlets.  Native  of  the  Himalaya  Mountains.  Several 
trees  are  around  the  Cactus  Garden. 

5.  Cupressus  Macnabiana  Murr.     MacNab  Cypress. 

Leaves  ovate,  obtuse,  thickened  at  apex,  conspicuously  glandular  and 
very  fragrant;  staminate  flowers  very  small,  globose,  with  1-2  lateral  anthers 
in  each  row;  cones  subglobose,  ^-^4  incn  broad;  scales  usually  6,  with  a 
prominent  conical  umbo;  seeds  reddish  brown,  with  a  broad  light-colored 
hilum. 

Low  spreading  tree  or  shrub,  or  sometimes  forming  a  pyramidal  tree 
20  feet  high  or  more.  Native  of  northern  California,  extending  from  Lake 
County  to  the  vicinity  of  Mt.  Shasta,  and  also  in  Lassen  County.  Several 
trees  are  between  the  Library  and  University  Avenue.  Flowers  are  pro- 
duced abundantly  almost  the  year  round. 

6.     Cupressus  funebris  Endl.     Funeral  Cypress. 

Branchlets  somewhat  flattened;  leaves  deltoid-ovate,  light  green;  cones 
short-peduncled,  globose,  l/$  inch  broad;  scales  8,  with  a  short  mucro. 

Tree  often  60  feet,  with  wide-spreading  branches  curved  upwards  and 
bearing  long  slender  pendulous  branchlets.  Native  of  China.  Several  trees 
are  near  the  Cactus  Garden  and  others  are  on  the  west  side  of  University 
Avenue  near  the  center  of  the  Arboretum. 

13.  Chamaecyparis. 

Evergreen  trees  with  flattened  branchlets  densely  clothed  with  opposite 
scale-like  leaves  in  4  rows.  Flowers  monoecious  on  separate  branchlets; 
staminate  oblong,  pistillate  subglobose.  Cones  globose,  maturing  the  first 


GROUP   OF    FIVE-LEAVED    PINES. 

Pinus   Lambertiana  Pinus    monticola 

(Reduced  to  one-third  natural  size.) 


Pinus    Strobus 
Pinus    excelsa 


GROUP    OF    THREE-LEAVED    PINES. 

Pinus    radiata  Pinus  canariensis 

Pinus   ponderosa 

Pinus    Coulteri  Pinus  Sabiniana 

(Reduced  to  one-third  natural  size.) 


GROUP    OF    TWO-LEAVED    PINES. 

Pinus  halepensis  Pinus  sylvestris  Pinus  Pinaster 

Pinus  Pinea  Pinus  nigra  Pinus  muricata 

(Reduced  to  two-fifths  natural  size.) 


•J^: 

fm 


/  yy 

: 


GROUP    OF    THE    ABIETE^E. 

Abies  cephalonica  Picea  excelsa 

Cedrus  Libani  Pseudotsuga   taxifolia 

(Reduced  to  one-half  natural  size.) 


GROUP    OF   THE    TAXODIEyE. 

Sequoia  sempervirens          .  Sequoia  Washingtoniana 

Cryptomeria  japonica  Taxodium  distichum 

(Reduced  to  one-half  natural  size.) 


GROUP  OF   THE   CUPRESSE/E. 


Cupressus  macrocarpa 
Chamsecyparis    Lawsoniana 

(Reduced  to  one-half  natural  size.) 


Thuya  orientalis 
Libocedrus  decurrens 


GYMNOSPERMS   ON    STANFORD   GROUNDS ABRAMS  109 

year;   scales  abruptly  dilated  and  flattened  at  apex,  with  short,  prominent 
points;  seeds  1-5  to  each  scale,  slightly  compressed. 

Six  species  are  known.  They  are  confined  to  the  Atlantic  and  Pacific 
Coast  regions  of  North  America  and  to  Japan  and  Formosa. 

1.     Chamaecyparis   Lawsoniana   Parl.     Lawson   Cypress. 

Leaves  closely  appressed  to  the  flattened  frond-like  branchlets,  bright 
green  and  with  a  gland  on  the  back,  paler  beneath  with  whitish  markings ; 
staminate  flowers  red;  cone  about  l/z  inch  broad,  red  brown. 

Tree  sometimes  200  feet  high,  with  horizontal  spreading  and  usually 
pendulous.  Native  of  the  coastal  region  of  Oregon  and  northern  Cali- 
fornia. A  variable  species  with  over  60  garden  forms.  Young  trees  are  along 
Alvarado  Row,  others  are  in  the  Arboretum  but  they  endure  our  dry  season 
badly.  A  handsome  specimen  is  in  the  lawn  at  the  Stanford  Residence  and 
another  is  at  9  Lasuen  Street. 

14.     Juniperus. 

Evergreen  trees  with  opposite  or  ternate,  scale-like  or  needle-shaped 
leaves.  Flowers  are  dioecious  or  monoecious,  minute ;  staminate  oblong-ovate ; 
anthers  with  4-8  pollen-sacs.  Cones  globose  or  oblong,  with  2-3  series  of 
fleshy  coalescent  scales,  berry-like,  maturing  the  second  year;  seeds  1 -several, 
ovate,  terete  or  angled. 

A  genus  of  about  30  species  widely  scattered  over  the  northern  hemi- 
sphere from  the  Arctic  Circle  to  the  mountains  of  the  subtropical  regions. 
Flowers  monoecious,  axillary;  leaves  jointed  at  base,  spreading  and  needle- 
shaped.  1.  /.  communis. 
Flowers  dioecious,  terminal ;  leaves  not  jointed  at  base,  often  scale-like. 
Leaves  all  alike,  in  3s,  slightly  spreading;  prostrate  shrub. 

2.  /.   recurva  squamata. 
Leaves  often  of  2  kinds,  those  on  vigorous  shoots  in  3s,  acicular,  those  on 

foliage  branches  in  2s,  minute  and  scale-like. 
Trees;  fruit  erect.  3.  /.  mrginiana. 

Low   shrub   with   procumbent  branches;    fruit  pendulous   on   recurved 
peduncles.  4.  /.  Sabina. 

1.     Juniperus  communis  L. 

Leaves  widely  spreading,  jointed  at  the  base,  narrowly  linear-lanceo- 
late, sharp-pointed,  l/2  inch  long  or  less,  concave  and  with  a  broad  white 
band  above;  fruit  sub-sessile,  dark  blue  and  glaucous,  %. -^  inch  broad. 

A  small  tree  or  erect  shrub.  Widely  scattered  over  the  northern  hemi- 
sphere. A  variable  species  with  many  geographical  varieties  and  garden 
forms.  One  specimen  is  west  of  the  Cactus  Garden. 


110  DUDLEY    MEMORIAL    VOLUME 

la.     Junipenis  communis  hemispherica  Parl. 

A  low  dense  shrub;  leaves  */j  inch  long  or  less,  stouter  and  stiffer  than 
those  of  the  typical  form. 

A  geographical  variety  found  in  the  mountains  of  southern  Europe  and 
northern  Africa.  One  bush  on  the  west  side  of  the  Cactus  Garden  near  the 
typical  form. 

Ib.     Juniperus   communis   oblonga    Loud. 

Erect  shrub  with  widely  spreading  branches  and  pendulous  branchlets; 
leaves  strongly  concave,  the  longest  ^4  mcn  l°ng- 

A  geographical  variety,  native  of  Transcaucasia.  One  specimen  is 
west  of  the  Cactus  Garden. 

2.     Juniperus  recurva  squamata  Parl. 

Leaves  in  3s,  straight,  slightly  spreading,  linear-lanceolate,  %  inch  long, 
sharp-pointed;  fruit  ^  inch  broad,  1 -seeded. 

Prostrate  shrub  with  long  trailing  branches.  Native  of  the  Himalaya 
Mountains.  One  specimen  is  on  the  southwest  border  of  the  Cactus  Garden. 

3.     Juniperus  virginiana  L.     Red  Cedar. 

Leaves  of  leading  shoots  acicular,  in  3s,  those  of  the  foliage  branches 
in  2s,  small  and  scale-like,  acute  or  acutish;  fruit  erect  on  short  peduncles, 
globose  or  ovoid,  y$  inch  broad,  1 -seeded. 

Tree  with  a  maximum  height  of  about  100  feet,  with  spreading  branches 
and  often  pendulous  branchlets.  A  variable  species  with  many  garden  forms. 
Native  of  eastern  North  America.  One  tree  with  wide  spreading  branches 
and  pendulous  branchlets  is  east  of  the  Cactus  Garden,  and  another  smaller 
and  more  compact  specimen  is  west  of  the  Cactus  Garden,  near  J.  communis. 

4.     Juniperus  Sabina  L. 

Leaves  of  leading  shoots  often  in  3s  and  acicular,  those  of  the  foliage 
branches  scale-like,  in  2s,  obtusish;  fruit  pendulous,  on  elongated  curved 
peduncles,  globose,  1-3-seeded. 

A  variable  species  with  many  garden  forms.  Native  of  eastern  North 
America,  Europe  and  Asia.  There  are  several  staminate  specimens  along 
the  border  of  the  Cactus  Garden. 


THE  SYNCHYTRIA  IN  THE  VICINITY  OF  STANFORD 
UNIVERSITY. 

By  JAMES  McMuRPHY,  Instructor  in  Botany. 

THE  GENUS  Synchytrium  belongs  to  the  Chytridiales,  the  lowest 
order  of  true  fungi.  The  majority  of  the  simple  forms  making 
up  this  order  are  parasitic  upon  protozoa,  algae  and  other  fungi,  but 
the  Synchytria  and  some  others  are  parasitic  upon  higher  plants.  All  forms 
yet  found  here  belong  to  the  sub-genus  Eusynchytrium,  in  which  both  resting 
spores  and  summer  sori  are  present.  And  it  is  the  so-called  summer  sorus 
that  is  meant  in  this  paper  when  the  word  sorus  is  used,  and  not  that  formed 
directly  by  the  germination  of  the  resting  spore.  There  are  no  sori  formed 
here  in  summer. 

Most  of  the  parasitic  fungi  are  supposed  to  attack  only  one  species  of 
host  or  to  be  limited  to  plants  nearly  related  botanically,  but  the  Synchytria 
are  not  supposed  to  be  so  limited.  The  hosts  given  for  S.  globosum  of 
Europe  are  "Violaceae,  Rosacese,  Compositae,  Rubiaceae,  etc.,"  plants  of 
widely  different  botanical  orders.  I  have  seen  no  account  of  any  Synchytrium 
having  been  transferred  from  one  host  to  another  under  laboratory  condi- 
tions. Since  the  species  of  Synchytrium  are  distinguished  by  the  presence  or 
absence  of  sori,  the  shape  and  general  appearance  of  the  galls  produced,  and 
the  shape  and  size  of  the  resting  spores,  it  may  be  asked  if  these  characters 
would  be  so  modified  by  a  change  of  host  as  to  make  the  fungus  unrecog- 
nizable. 

In  April,  1910,  I  found  near  Stanford  University  what  appeared  to  be 
a  Synchytrium  growing  on  Amsinckia  intermedia  and  took  a  small  quantity 
to  the  laboratory,  where  I  found  the  material  to  contain  both  sori  and  resting 
spores.  Some  days  later  I  gathered  more  material  and  found  that  only  rest- 
ing spores  were  present,  sori  not  having  been  formed  during  the  ctear,  warm 
days  preceding.  At  the  same  time  I  found  growing  with  the  Amsinckia, 
Erodium  cicutarium  bearing  S.  papillatum  with  only  resting  spores.  Now  S. 
papillatum  is  known  only  from  California  and  is  supposed  to  be  indigenous 
here,  though  it  is  found  only  on  the  above  named  host,  which  is  a  weed  intro- 
duced from  Europe. 

As  there  were  no  more  long  periods  of  wet  weather,  I  was  unable  to  get 
any  zoospores  for  cross  infections,  but  in  March  of  the  following  year  I  had 
an  abundance  of  zoospores  from  both  forms.  I  was  unable  to  infect  Am- 
sinckia with  zoospores  from  Erodium  or  to  infect  Erodium  with  those  from 


112  DUDLEY    MEMORIAL   VOLUME 

Amsinckia.  In  March  of  this  year,  I  placed  an  inch  or  so  of  the  ends  of  the 
leaves  of  a  few  potted  Erodiums  and  Amsinckias  into  a  vessel  of  water  con- 
taining zoospores  of  the  Synchytrium  on  Amsinckia.  After  about  two  hours 
the  plants  were  removed  to  a  damp  place.  Some  of  the  leaves  of  Amsinckia 
developed  an  abundance  of  Synchytrium  on  the  ends  that  had  been  immersed 
in  the  water,  but  there  was  none  on  the  Erodium.  I  have  had  no  zoospores 
for  further  experiment. 

In  the  hills  I  have  found  Erodium  cicutarium  growing  with  Erodium 
botrys,  nearly  every  plant  of  the  former  more  or  less  discolored  by  Synchyt- 
rium papillatum,  but  the  latter  entirely  free  of  the  parasite.  Near  the 
Museum  and  the  Nursery,  I  found  Erodium  cicutarium  growing  with  Ero- 
dium moschatum,  the  one  often  discolored  with  Synchytrium,  the  other  with 
none.  These  two  species  of  Erodium  are  so  similar  in  appearance  that  be- 
ginners in  botany  often  find  it  difficult  to  distinguish  them,  but  the  Synchyt- 
rium zoospores  seem  to  have  no  such  difficulty. 

If  Synchytrium  papillatum  is  really  endemic  on  some  native  California 
plant,  then,  since  we  have  no  native  Erodiums,  it  must  have  passed  to  E. 
cicutarium  from  some  host  much  farther  removed  botanically  than  either  of 
the  two  mentioned  above. 

The  following  species  of  Synchytrium  have  been  found  in  the  vicinity 
of  Stanford  University: 

SYNCHYTRIUM    PAPILLATUM    Farlow. 

Synchytrium  papillatum  Farlow.  Bull.  Bussey  Inst.  2  :239.  Bot.  Gaz. 
10:239,  1885. 

"Spots  dark  purple,  galls  glandular,  formed  of  papillate,  pyriformly 
swollen  epidermal  cells,  resting  spores  elliptical,  .06-.07  mm.  by  .04-.05  mm., 
epispore  brown,  somewhat  roughened.  Sori  superficial,  spherical,  .10-.  12  mm. 
in  diameter." 

"On  leaves  of  Erodium  cicutarium  L'Her.     California." 

In  late  winter  and  early  spring  this  species  may  be  found  on  the  slopes 
of  the  hills  as  well  as  in  the  valley,  and  often  there  are  places  where  it  is 
difficult  to  find  plants  of  the  Red-stemmed  Filaree  that  are  entirely  free  of  the 
deep  red  (dark  purple  when  dried)  discolorations  caused  by  the  parasite. 

The  galls  vary  considerably  in  size  and  may  be  nearly  regular  in  outline 
or  irregular  and  strongly  papillate.  Usually  there  are  only  one  or  two  rest- 
ing spores  in  a  gall,  but  there  are  sometimes  four  or  five.  The  sori  are 
spherical,  flattened  or  elongated  and  contain  about  45-70  zoosporangia 
15-21  x  15-26/i  in  diameter.  In  material  taken  after  a  week  or  two  of  rainy 
weather  I  have  found  sori  more  abundant,  but  after  a  few  days  of  sunshine 
only  resting  spores. 


SYNCHYTRIA   IN   VICINITY  OF    STANFORD MC  MURPHY  113 

SYNCHYTRIUM  AMSINCKI^E    n.  sp. 

Spots  light  yellow  when  immature,  reddish  brown  at  maturity;  galls 
formed  of  swollen,  externally  projecting  epidermal  cells,  which  collapse 
when  mature ;  resting  spores  one  or  two  in  a  cell,  elliptical  or  globose, 
70-100  x  90-11 5/x,  epispore  brown,  smooth;  sorus  yellow,  elliptical  or  glo- 
bose, in  galls  similar  to  those  of  the  resting  spores,  65-90  x  100-120/x,;  zoo- 
sporangia  in  a  sorus,  7-20,  angular  from  mutual  pressure,  or  globose  to 
elliptical,  25-40  x  25-45/i  in  diameter. 

On  leaves  and  stems  of  Amsinckia  intermedia  F.  &  M.  Stanford  Uni- 
versity, California,  in  fields  and  waysides. 

This  is  closely  related  to  the  European  species  S.  Myosotides  Kuhn,  but 
that  species,  so  far  as  I  know,  has  never  been  known  to  produce  sori,  and 
belongs  to  the  sub-genus  Pycnochytrium,  while  in  this  form  sori  are  common 
and  the  galls,  which  are  often  200-26 5/u,  in  diameter  before  collapsing  at 
maturity,  are  brown  rather  than  deep  red,  as  in  that  species. 

The  type  was  collected  near  Stanford  University  March  24,  1911,  and 
is  deposited  in  the  Dudley  Herbarium. 

SYNCHYTRIUM  INNOMINATUM  Farlow. 

Synchytrium  innominatum  Farlow.     Bot.  Gaz.  10:240,  1885. 

"Spots  dark  red,  resting  spores  globose  or  slightly  elliptical,  .07-.  10  mm. 
in  diameter,  epispore  thin  and  smooth,  in  oval  host  cells,  which  do  not  pro- 
ject beyond  the  surface  of  the  leaves.  Sori  yellow,  about  .12-.  15  mm.  in 
diameter,  sunk  in  the  leaves." 

"On  leaves  of  Malacothrix.     Santa  Cruz,  Cal." 

On  leaves  of  Agoseris  grandiflora  Greene,  Page  Mill  Road,  altitude 
about  1,000  feet. 

My  specimens  are  referred  to  the  above  species,  though  the  resting  spores, 
of  which  there  are  one  or  two  in  a  cell,  are  more  often  elliptical  than  globose 
and  vary  more  in  size,  40-80  x  60-90/A.  The  sori  are  also  smaller, 
70-117  x  78-117)Lt.  As  Dr.  Farlow  says,  1.  c.,  "the  species  is  certainly  closely 
related  to  S.  Taraxici  D.  By.  &  Wor."  But  the  sori  in  my  material  are 
larger  on  the  average  and  the  discolorations  on  the  leaves,  when  fresh, 
were  dark  red  rather  than  golden  red  or  blood  red. 

It  is  very  desirable  that  cultures  be  made  on  the  dandelion  and  related 
plants,  that  the  limits  of  the  species  may  be  determined. 

SYNCHYTRIUM    ANDINUM    Lagh. 

Synchytrium  andinum  Lagh.     Bull.  Boiss.    1895,  p.  61. 
Galls   multicellular,    reddish   brown,    often   confluent;    sori   globose   or 


114  DUDLEY    MEMORIAL    VOLUME 

elliptical,  60-110  in  diameter;  zoosporangia  numerous,  irregular,  angular 
from  mutual  pressure,  40-60  in  diameter,  contents  golden  yellow;  resting 
spores  solitary,  80-120  in  diameter,  epispore  thick,  smooth,  dark  brown. 

On  leaves  of  Ranunculus,  Quito,  Ecuador. 

On  stems,  leaves  and  floral  parts  of  Ranunculus  calif  ornicus  Benth,  wet 
ground  between  Stanford  University  and  Mayfield. 

In  the  spring  of  1911,  many  of  the  buttercups  were  badly  deformed  by 
this  fungus,  the  ovaries,  stamens  and  petals  being  sometimes  attacked  as  well 
as  the  leaves  and  stems,  but  this  spring,  presumably  on  account  of  the  very 
light  rainfall,  I  was  unable  to  find  more  than  half  a  dozen  leaves  attacked 
by  the  parasite. 

My  specimens,  while  agreeing  very  well  with  the  above  description  taken 
from  Saccardo,  show  somewhat  greater  variation  in  size  of  resting  spores 
and  sori,  and  the  zoosporangia  are  considerably  smaller,  26-34  x  26-48,  usu- 
ally 15-30  in  a  sorus.  The  galls,  when  solitary,  are  hemispherical  or  sub- 
globose  and  150-250  in  height. 

UROPHLYCTIS   PLURIANNULATUS    Farlow. 
Urophlyctis   pluriannulatus    Farlow,    Rhodora    10:13.     1908. 
Uromyces  pluriannulatus  B.  &  C.,  Grevillia  3:57.    1874. 
Synchytrium  pluriannulatum  Farlow,   Bot.   Gaz.   10:243.    1885. 
This  fungus  was  found  growing  on  Saniclua  Menziesii  H.  &  A.  along 
the  creek  below  Searsville  Lake. 


EXPLANATION    OF   PLATE. 


Fig.  1.      Synchytrium  papillattim,  sorus  and  two  galls  with  resting  spores,  about  125  diameters. 

Fig.  2-3.  5.  Amsinckiae,  sections  through  sorus  and  three  galls  with  resting  spores,  about  140  diameters. 

Fig.  4.      S.  Amsinckiae,  zoospores,  from  stained  preparation,  500  diameters. 

Fig.  5.      5.  innominalum,  section  through  sorus  and  resting  spore,  about  150  diameters 

Fig.  6-7.  5.  Andinum,  sections  through  sorus  and  two  resting  spores,  about  120  diameters. 


Fig.  1.  Agoseris  grandiflora,  with  very  light 
attack  of  Synchytrium  innominatum. 

Fig.  2.  Ranunculus  californicus  with  Syn- 
chytrium andinum.  The  leaves  and  floral  parts 
of  the  plant  at  the  right  are  much  deformed. 


Fig.  3.  Amsinckia  intermedia,  deformed  by 
Synchytrium  Amsinckiae. 

Fig.  4.  Erodium  cicutarium,  with  parts  of  the 
leaves  and  petioles  darkened  by  Synchytrium 
papillatum. 


THE  LAW  OF  GEMINATE  SPECIES.* 
By  DAVID  STARR  JORDAN,  President  of  Stanford  University. 

IN  "Evolution  and  Animal  Life,"  by  Jordan  and  Kellogg  (page  120),  the 
following  words  are  used: 
"Given  any  species,  in  any  region,  the  nearest  related  species  is  not  to 
be  found  in  the  same  region  nor  in  a  remote  region,  but  in  a  neighboring 
district  separated  from  the  first  by  a  barrier  of  some  sort  or  at  least  by  a  belt 
of  country,  the  breadth  of  which  gives  the  effect  of  a  barrier." 

Substituting  the  word  "kind"  for  species  in  the  above  sentence,  thus' 
including  geographical  subspecies,  or  nascent  species,  as  well  as  species 
clearly  definable  as  such,  Dr.  J.  A.  Allen  accepts  this  proposition  as  repre- 
senting a  general  fact  in  the  relations  of  the  higher  animals.  To  this  gen- 
eralization Dr.  Allen,  in  Science,  has  given  the  name  of  "Jordan's  Law." 
The  present  writer  makes  no  claim  to  the  discovery  of  this  law.  The  lan- 
guage above  quoted  is  his,  but  the  idea  is  familiar  to  all  students  of  geo- 
graphical distribution  and  goes  back  to  the  master  in  that  field,  Moritz 
Wagner.  Dr.  Wagner  was  one  of  the  most  clear-sighted  and  long-headed  men 
of  the  early  evolutionists.  In  recognizing  the  potency  of  isolation  in  species- 
forming  he  made  the  mistake,  however,  of  not  recognizing  selection  as  the 
basis  of  adaptation. 

This  law  rests  on  the  fact  that  the  minor  differences  which  separate 
species  and  subspecies  among  animals  are  due  to  some  form  of  segregation 
or  isolation.  Selection  produces  adaptation,  but  the  distinctive  characters 
of  species  are  in  general  non-adaptive.  They  find  their  origin  in  the  different 
currents  of  life  which  isolation  makes  possible.  By  some  barrier  or  other 
the  members  of  one  group  are  prevented  from  interbreeding  with  those  of 
another  minor  group  or  with  the  mass  of  the  species.  As  a  result,  local 
peculiarities  arise.  "Migration  holds  species  true,  localization  lets  them 
slip,"  or  rather  leaves  them  behind  in  the  process  of  modification.  The 
peculiarities  of  the  parents  in  an  isolated  group  become  intensified  by  in  and 
in  breeding.  They  become  modified  in  a  continuous  direction  by  the  selec- 
tion induced  by  the  local  environment.  They  are  possibly  changed  in  one 
way  or  another  by  germinal  reactions  from  impact  of  environment.  At  last 
a  new  form  is  recognizable.  And  this  new  form  is  never  coincident  in  its 
range  with  the  parent  species,  or  with  any  other  closely  cognate  form,  neither 
is  it  likely  to  be  located  in  some  remote  part  of  the  earth.  Whenever  the 
range  of  two  such  forms  overlaps  in  any  degree,  the  fact  seems  to  find  an 


*  Most  of  this  paper  was  published  in  the  American  Naturalist,  vol.  42,  pp.  73-80, 
February,  1908,  under  the  same  title. 


116  DUDLEY    MEMORIAL    VOLUME 

explanation  in  reinvasion  on  the  part  of  one  or  both  of  the  forms.  The 
obvious  immediate  element  in  the  formation  of  species  is,  therefore,  isolation, 
and  behind  this  are  the  factors  of  heredity,  of  variation,  of  selection,  and 
others  as  yet  more  or  less  hypothetical.  The  formation,  through  segregation, 
of  different  breeds  of  sheep  in  the  different  countries  of  England,  as  noted  by 
Jordan  and  Kellogg  (p.  82),  seems  exactly  parallel  with  the  formation  of 
species  in  nature.  In  like  manner,  the  occasional  development  of  breeds 
arising  from  the  peculiarities  of  individuals  is  possibly  parallel  with  the 
"mutations"  of  the  evening  primrose.  It  seems  to  me  probable,  however, 
that  these  mutations  are  phenomena  of  hybridism.  Such  breeds  are  the 
Ancon  sheep  in  Connecticut  and  the  blue-cap  Wensleydale  sheep  in  Aus- 
tralia. The  hornless  Hereford  cattle  lately  established  in  Kansas  is  a  case 
in  point.  The  "ontogenetic  species" — groups  in  which  many  individuals  are 
simultaneously  modified  in  the  same  way  by  like  conditions  of  food  or  cli- 
mate— show  no  permanence  in  heredity.  Such  forms,  however  strongly 
marked,  should,  therefore,  have  no  permanent  place  in  taxonomy.  The  recent 
studies  of  Mr.  Beebe  on  the  effects  of  moist  air  in  giving  dusky  colors  to 
birds  serve  to  illustrate  the  impermanence  of  the  groups  or  subspecies  char- 
acterized by  dark  shades  of  color  developed  in  regions  of  heavy  rainfall. 
These  dark  shades  are  not  inherited  and  are  not  constant  in  the  same  in- 
dividual if  it  is  brought  under  new  conditions. 

It  may  also  be  noted  in  passing  that  one  cause  of  the  potency  of  arti- 
ficial selection  among  domesticated  animals  or  cultivated  plants  is  that  such 
selection  is  always  accompanied  by  segregation.  The  latter  is  taken  for 
granted  in  discussions  of  this  topic  and  hence  its  existence  as  a  factor  is 
usually  overlooked.  While  poultry  or  pigeons  can  be  rapidly  and  radically 
changed  by  artificial  selection,  in  isolation,  no  process  of  selection  without 
isolation  can  have  any  permanent  result.  For  example,  we  know  no  way 
of  improving  the  breed  of  salmon,  because  the  salmon  we  have  selected  for 
reproduction  must  be  turned  loose  in  the  sea,  where  they  are  at  once  lost 
in  the  mass. 

New  forms  of  gold  fish,  carp  and  other  domesticated  fishes  can  be  made 
easily  by  selection,  because  these  fishes  can  be  kept  in  aquaria  or  in  little 
ponds,  but  new  forms  of  mackerel  or  herring  are  beyond  the  control  of  man 
and  the  species  actually  existing  have  been  of  the  slowest  creation,  their 
origin  lost  in  geologic  times. 

One  of  the  most  interesting  features  of  "Jordan's  law"  is  the  existence 
of  what  I  have  termed  geminate  species — twin  species — each  one  representing 
the  other  on  opposite  sides  of  some  form  of  barrier.  In  a  general  way,  these 
geminate  species  agree  with  each  other  in  all  the  respects  which  usually  dis- 
tinguish species  within  the  same  genus.  In  all  matters  of  selection  and  adap- 


LAW    OF    GEMINATE    SPECIES JORDAN  117 

tation  they  are  absolutely  identical.  They  are  usually  identical  in  habits. 
They  differ  in  minor  regards,  characters  which  we  may  safely  suppose  to  be 
of  later  origin  than  the  ordinary  specific  characters  in  their  group.  Illus- 
trations of  geminate  species  of  birds,  of  mammals,  of  fishes,  of  reptiles,  of 
snails,  or  of  insects,  are  well  known  to  all  students  of  these  groups,  and 
illusrations  may  be  found  at  every  hand. 

Each  island  of  the  West  Indies,  which  is  well  separated  from  its  neigh- 
bors, has  its  own  form  (species  or  subspecies)  of  golden  warbler  (Dendroica 
cestiva).  Each  island  in  the  South  Seas  has  its  geminate  forms  of  reptiles 
or  fishes.  Each  island  of  the  Hawaiian  group  has  its  own  representative  of 
each  one  of  the  types  or  genera  of  Drepanida.  Each  of  the  three  groups  of 
rookeries  in  Bering  Sea  has  its  own  species  of  fur  seal.  Callorhinus  alascanus 
on  the  Pribilof  Islands,  C.  ursinus  on  the  Commander  Islands,  and  C.  curi- 
lensis  of  the  Kuriles  and  Robben  Islands,  these  species  most  clearly  related 
but  showing  no  intergradations,  because  no  intercrossing  is  possible,  each 
having  its  own  distinct  line  of  migrations. 

Similar  conditions  exist  among  species  of  fresh  water  fishes,  the  world 
over.  Dr.  Ortmann  has  described  the  conditions  of  species  forming  through 
isolation  in  the  river  cray  fish  (Cambarus)  ;  Dr.  Gulick  in  the  land  snails 
(Helix)  and  various  writers  in  the  river  mussels  (Unionidce). 

Most  clearly  marked  cases  of  geminate  species  occur  among  the  fishes 
on  the  two  sides  of  the  isthmus  of  Panama.  Living  under  essentially  the 
same  conditions,  but  separated  since  the  end  of  the  Miocene  Period  by  the 
rise  of  the  isthmus,  we  find  species  after  species  which  has  been  thus  split 
into  two.  These  geminate  species,  a  hundred  or  more  pairs  in  number,  were 
at  first  regarded  as  identical  on  the  two  shores  of  the  isthmus.  Later  one 
pair  after  another  was  split  into  recognizable  species.  The  latest  authority 
on  the  subject,  Mr.  C.  T.  Regan,  seems  to  doubt  if  any  species  of  shore  fishes 
are  actually  identical  on  the  two  sides  of  the  isthmus. 

To  make  this  clear,  though  at  the  risk  of  being  tedious,  I  give  below  a 
partial  list  of  these  genuine  species  about  the  isthmus  of  Panama: 

Atlantic    Coast  Pacific    Coast 

Harengula  humeralis  Harengula  thrissina 

Clupanodon  oglinus  Clupanodon  libertatis 

Centropomus  undecimalis  Centropomus  viridis 

Centropomus  pedimacula  Centropomus  medius 

Centropomus  affinis  Centropomus  ensiferus 

Epinephelus  adscensionis  Epinephelus  analogus 

Alphestes  afer  Alphestes  multiguttatus 

Dermatolepis  inermis  Dermatolepis  punctatus 

Hypoplectrus   unicolor  Hypoplectrus  lamprurus 


118  DUDLEY    MEMORIAL   VOLUME 

Atlantic    Coast  Pacific    Coast 

Lutianus  cyanopterus  Lutianus  novemfasciatus 

Lutianus   apodus  Lutianus  argentiventris 

Lutianus  analis  Lutianus  Colorado 

Lutianus  synagris  Lutianus  guttatus 

Hcemulon  album  Hcemulon   sexfasdatum 

Hcemulon  parra  Hcemulon  scudderi 

Hcemulon  schrancki  Hcemulon  steindachneri 

Anisotremus  surinamensis  Anisotremus  interruptus 

Anisotremus  virginicus  Anisotremus  toeniatus 

Conodon  nobilis  Conodon  serrifer 

Pomadasis  crocro  Pomadasis  branicki 

Calamus  macrops  Calamus  taurinus 

Xystcema   cinereum  Xystozma  simillimum 

Eucinostomus  pseudogula  Eucinostomus  dowi 

Kyphosus  incisor  Kyphosus  analogus 

Isopisthus  parvipinnis  Isopisthus  remifer 

Nebris  microps  Nebris  zestus 

Larimus  fasciatus  Larimus  pacincus 

Odontoscion  dentex  Odontoscion  xanthops 

Corvula  sialis  Corvula  macrops 

Bairdiella  verce-crucis  Bairdiella  armata 

Micropogon  furnieri  Micropogon  ectenes 

Umbrina  broussoneti  Umbrina  xanti 

Menticirrhus  littoralis  Menticirrhus  elongatus 

Eques  acuminatus  Eques  viola 

This  list  may  be  greatly  extended,  but  the  series  noted  will  illustrate  the 
point  in  question.  Whenever  a  distinct  and  sharply  denned  barrier  exists, 
geminate  or  twin  species  may  be  found  on  the  two  sides  of  it,  unless,  as  some- 
times happens,  the  species  has  failed  to  maintain  itself  on  one  side  or  the 
other  of  the  barrier.  So  far  as  Panama  is  concerned,  we  have  evidence 
that  the  barrier  was  raised  near  the  end  of  Miocene  time  with  no  trace  of 
subsequent  depression.  We  can  thus  form  some  estimate  of  the  age  of 
separation  in  at  least  a  small  number  of  closely  related  species.  In  this  and 
similar  cases  it  is  not  possible  to  conceive  of  the  formation  of  these  species 
by  sudden  mutation,  or  that  they  would  retain  their  separate  existence  were 
the  element  of  segregation  removed.  While  segregation  or  isolation  is  not  a 
force,  and  perhaps  not  strictly  a  cause  in  species  formation,  it  is  a  factor 
which  apparently  can  never  be  absent,  if  the  species  retains  its  independent 
existence. 

There  is  no  doubt  that  the  distribution  of  higher  animals  in  general  is  in 
accord  with  "Jordan's  law."  Examples  by  the  thousand  come  up  from  every 
hand.  If  we  had  a  hundredth  part  of  the  amount  of  available  evidence  in 
support  of  mutation  theories,  these  theories  would  pass  from  the  realm  of 


LAW    OF    GEMINATE    SPECIES JORDAN  119 

hypothesis  into  that  of  fact.  But  the  application  of  this  law  or  rule  to  plants 
and  to  one-celled  animals  has  been  questioned.  So  far  as  rhizopods  are  con- 
cerned, Dr.  Kofoid  finds  that  the  species  are  in  general  sharply  defined  and 
of  the  widest  distribution  in  the  sea,  so  that  we  can  hardly  state  laws  as  de- 
fining their  geographical  distribution.  To  these  minute  floating  animals,  the 
sea  scarcely  offers  barriers  at  all,  and  the  recognized  species  do  not  seem  to  be 
products  of  geographical  isolation.  Doubtless  these  species  in  duration  and 
in  nature  correspond  more  nearly  to  genera  or  families  of  higher  animals 
than  to  actual  species.  Perhaps  minor  specific  differences  such  as  we  note 
among  arthopods  or  vertebrates  are  intangible  or  non-existent.  The  effects 
of  isolation  may  be  tangible  only  among  forms  which  possess  more  varied 
relations  with  their  environment. 

The  application  of  this  law  to  plants  has  also  been  denied.  But  gem- 
inate species  are  just  as  common  in  botany  as  in  zoology,  and  the  effects 
of  isolation  in  species-forming  are  just  as  distinct.  The  law  is  just  as  patent 
in  the  one  case  as  in  the  other.  It  is  merely  obscured  by  other  laws  or  con- 
ditions which  obtain  among  plants. 

In  the  nature  of  things,  most  physical  barriers  are  more  easily  crossed 
by  plants  than  by  animals.  The  possibilities  of  reinvasion  are  thus  doubtless 
much  increased.  The  plant  is  limited  by  climate,  rainfall,  nature  of  soil,  and 
the  same  species  is  likely  to  occupy  all  suitable  locations  within  a  large  area. 
Animals  are  more  mobile  than  plants  within  their  range,  a  fact  which  tends 
to  keep  the  interbreeding  masses  more  uniform.  In  the*  struggle  for  exist- 
ence, the  plant  is  pitted  against  its  environment.  Whether  the  plant  survives 
or  not  depends  not  much  on  the  nature  of  the  seed,  but  mainly  on  its  relation 
to  the  spot  on  which  it  falls.  There  is  little  selection  within  the  species  due 
to  the  choice  of  one  individual  as  against  another.  Selection  can  only  happen 
where  plants  are  overcrowded,  and  there  the  survival  is  mainly  that  of  the 
seed  whose  roots  run  deepest.  There  is  little  room  for  struggle  between 
closely  related  species.  Each  individual  grows — if  it  can — on  the  spot 
where  it  falls.  The  variations  among  plants  are  great,  but  these  variations 
are  mostly  lost  unless  reinforced  by  segregation.  There  is  no  likelihood  of 
the  survival  of  DeVries'  mutants  (or  hybrids)  of  the  evening  primrose  if 
these  forms  are  left  free  to  mix  in  the  same  field. 

Among  plants  we  often  notice  the  fact — rare,  though  not  unknown 
among  animals — of  numerous  species  of  the  same  genus  occupying  the  same 
area.  In  some  cases  these  species  are  closely  related,  suggesting  mutants,  and 
in  other  cases  the  relation  indicates  the  existence  of  hybrids.  In  California, 
for  example,  there  are  in  the  same  general  region  many  species  of  Lupinus, 
of  Calochortus,  of  Ceanothus,  of  Arctostaphylos,  of  Eschscholtzia,  of 


120  DUDLEY    MEMORIAL   VOLUME 

Godetia,  of  QEnothera,  and  Opuntia.  Eucalyptus,  Acacia  and  Epacris  in 
Australia  are  examples  even  more  striking.  But  I  have  never  seen  very 
closely  related  or  geminate  forms  in  any  of  these  genera  actually  growing 
together.  I  suspect  that  they  do  so  sometimes  and  that  the  explanation  is 
found  in  reinvasion.  Dr.  G.  H.  Shull  of  the  Carnegie  Station  for  Experi- 
mental Evolution  tells  me  that  most  of  these  plants  are  self-fertilized,  a  con- 
dition unfavorable  to  Panmixia  or  the  loss  of  the  individual  or  local  varia- 
tion in  the  mass.  Self -fertilized  plants  may  be  neighbors  without  really 
"growing  together."  But  "growing  together"  is  an  indefinite  statement  as 
applied  to  plants.  The  elder,  the  alder  and  the  madrono  (Arbutus)  abound 
in  the  Santa  Clara  Valley.  But  no  one  ever  saw  any  two  of  these  trees  stand- 
ing side  by  side.  Each  has  its  limitations,  as  to  soil  and  moisture,  and  its 
own  choice  of  locations. 

Setting  aside  these  genera  which  are  represented  by  many  species  in  a 
limited  area,  and  among  which  mutation,  hybridism  and  self-fertilization  may 
be  conceivable  factors  in  species-forming,  we  find  the  law  of  geminate  species 
applying  to  plants  as  well  as  to  animals.  Crossing  the  temperate  zone  any- 
where on  east  and  west  lines,  we  find  species  after  species  replaced  across 
the  barriers  by  closely  related  forms.  Illustrations  may  be  taken  anywhere 
among  the  higher  plants — equally  well,  no  doubt,  among  lower  ones.  Many 
genera  are  local  in  their  distribution,  monotypic — with  a  single  species,  the 
origin  of  which  cannot  be  traced.  Such  species  spread  far  and  wide  without 
visible  change  within  the  species.  But  many  other  genera  belt  the  earth  or 
come  very  near  doing  so,  each  form  or  species  being  geminate  as  related  to 
its  next  neighbor.  This  fact  is  illustrated  in  Rubus,  Alnus,  Sambucus,  Pla- 
tanus,  Fagus,  Veratrum,  Symplocarpus,  Symphoricarpus,  Castanea,  Quercus, 
Pinus,  Tsuga,  Acer,  Rhus,  Pyrus,  Prunus,  Lonicera,  Ranunculus,  Trientalis, 
Lilium,  Trillium,  Veronica,  Aquilegia,  Gentiana,  Viola,  Epilobium,  Pteris, 
Mimulus,  Trifolium,  Solidago,  Aster,  Helianthemum,  Triosteum,  Geranium, 
Ribes,  Asarum,  Habenaria,  Saxifraga,  Clintonia,  Calycanthus,  Fraxinus, 
Philadelphus,  Crataegus,  Azalea,  Erythronium,  Rhododendron,  Viburnum, 
Cornus,  Cercis,  Eupatorium.  All  these  genera  and  many  others  furnish  an 
abundance  of  examples.  It  would  be  hard  to  find  a  widely-distributed  genus 
which  did  not. 

Taking  a  single  example,  the  pink-flowering  raspberry  of  the  eastern 
United  States,  Rubus  odoratus,  becomes  on  the  Pacific  Slope  the  white-flow- 
ered Rubus  parviflorus ,  ("  nutkanus" ) .  On  the  California  sea  coast,  Rubus 
velutinus,  with  tasteless  fruit,  again  takes  the  place  of  the  latter.  The  black 
raspberry,  Rubus  occidentalis,  is  replaced  westward  by  its  double,  Rubus  leu- 
codermis.  The  common  blackberry,  Rubus  mllosus,  is  replaced  in  the  eastern 


LAW    OF    GEMINATE    SPECIES JORDAN  121 

hills  by  Rubus  alle ghaniensis  and  in  the  far  west  by  Rubus  mtijolius,  while 
still  other  species  surround  the  world,  taking  its  place  in  Europe  and  in  Asia. 

We  may,  therefore,  say  that  with  plants  as  well  as  animals  geminate 
species  as  above  denned  owe  their  distinctness  to  some  form  of  isolation  or 
segregation,  and  that,  broadly  speaking,  with  occasional  exceptions,  given 
any  form  of  animal  or  plant  in  any  region,  the  nearest  related  form  is  not 
to  be  found  in  the  same  region  nor  in  a  remote  region,  but  in  a  neighboring 
region,  separated  from  the  first  by  a  barrier  of  some  sort,  not  freely  travers- 
able. 

A  law,  that  is,  an  observed  relation  of  cause  and  effect,  is  not  invalidated 
by  the  presence  of  other  effects  due  to  other  causes  in  the  same  environment. 
The  actual  conditions  in  nature  are  everywhere  not  products  of  single  and 
simple  forces,  but  resultants  of  many  causative  influences,  often  operative 
through  the  long  course  of  ages.  As  a  rule,  also,  related  species  in  almost 
every  group  are  connected  by  a  fringe  of  intergradations  we  call  subspecies. 
If  barriers  are  sharply  defined  geminate  species  are  sharply  defined,  also. 
If  barriers  are  diffuse,  we  find  geographical  subspecies  connecting  them, 
either  wholly  or  in  part.  There  is  no  difference  between  a  subspecies  and  a 
true  species  except  that  which  is  involved  in  sharpness  of  definition.  If  the 
barrier  cannot  be  crossed,  the  species  dependent  on  the  barrier  is  well  defined 
and  therefore  unquestionable,  however  small  the  elements  of  difference.  A 
subspecies,  if  real,  is  always  based  on  some  matter  in  geographical  distribu- 
tion. 

It  may  be  urged  that  these  geminate  groups  or  forms  are  not  true  species 
because  they  often  intergrade  one  into  another,  and  they  would  probably  be 
lost  by  intermingling  if  the  barriers  were  removed.  It  is  .sometimes  claimed 
that  only  physiological  tests  of  species  can  be  trusted,  as  true  species  will  not 
blend  and  their  hybrids,  if  formed,  will  be  sterile.  All  this  is  purely  hypo- 
thetical and  impracticable  to  the  systematic  zoologist,  and  not  of  much  value 
to  the  botanist.  Interbreeding  is  no  test  of  species.  Closely  related  species 
in  almost  any  group  of  plants  or  animals  can  usually  be  readily  crossed.  As 
the  relation  becomes  less  close,  partial  sterility  of  all  grades  and  then  total 
sterility  appear. 

If  the  term  species  has  any  meaning  at  all,  those  species  we  find  in  nature 
are  real  species.  Nothing  can  be  more  real  than  that  which  actually  exists. 
And  real  species  have,  as  a  rule,  indefinite  boundaries,  shading  off  into  sub- 
species, geminate  species,  ontogenetic  forms  and  the  like.  In  these  eccen- 
tricities we  must  humor  them.  As  Darwin  observed,  these  peculiarities  are 
fascinating  to  us  "as  speculatists"  however  "odious"  they  are  to  us  "as  sys- 
tematists."  And  if  we  are  to  understand  the  significance  of  nature,  we  have 


122  DUDLEY    MEMORIAL   VOLUME 

to  describe  these  facts  and  relations  as  they  actually  are.  Then  we  have  to 
find  out  what  changes  we  can  work  in  individuals  and  species  by  such  altera- 
tions of  conditions  as  experiment  can  give. 

We  do  not  know  actually  any  species  of  animal  or  plant  until  we  know 
all  changes  that  would  or  could  take  place  in  its  individuals  under  all  condi- 
tions of  environment. 

P.  S. — Since  this  was  written  I  have  received  the  charming  sketch  of 
Linnaeus  by  Professor  Edward  Lee  Greene.  In  a  discussion  of  Linnaeus 
as  an  evolutionist  Mr.  Greene  shows  very  clearly  that  the  great  botanist 
had  in  mind  the  same  fact  as  to  the  relation  of  species  which  I  have  in- 
dicated in  this  paper.  In  a  note  on  Thallctrum  lucidum  referring  to  the 
geminate  form,  he  says,  "This  plant  is  possibly  not  very  distinct  from 
T.  flavum.  It  seems  to  me  to  be  the  product  of  its  environment."  Of  these 
species  of  meadow  rue,  flavum  belongs  to  the  cool  moist  meadows  of  northern 
Europe,  and  lucidum  to  southern  France  and  Spain. 

Referring  again  to  the  seaside  Virgin's  Bower,  Clematis  maritima,  he 
says,  "Magnol  and  also  Ray  have  adjudged  this  to  be  a  variety  of  C.  flam- 
mula.  I  should  rather  think  it  is  derived  from  C.  recta  under  altered 
conditions." 

As  to  the  Siberian  yarrow,  as  compared  to  the  common  yarrow  of 
Europe,  he  says,  "May  not  the  Siberian  mountain  soil  and  climate  have 
moulded  this  out  of  A.  ptarmica?" 

Comparing  two  species  of  Kosteletzkya  (Hibiscus),  one  from  the  Adri- 
atic region,  the  other  from  the  salt  marshes  of  our  southern  states,  he  asks, 
"May  not  the  Venetian  species  have  sprung  from  the  Virginian?" 

Dr.  Greene  mentions  other  cases  of  a  similar  sort,  but  these  will  show 
that  the  idea  of  geminate  species  split  off  from  the  parent  by  separation  and 
changed  conditions  was  present  with  the  great  botanist. 


SOME  RELATIONS  BETWEEN  SALT  PLANTS  AND  SALT-SPOTS. 
By  WILLIAM  AUSTIN  CANNON,  Desert  Laboratory,  Tucson,  Arizona. 

ONE  of  the  characteristic  environmental  factors  which  desert  plants 
must  successfully  meet  is  the  high  salt  content  of  the  soils.  This  is 
especially  true  of  such  areas  as  have  poor  surface  drainage  and  where 
water  is  removed  only  by  evaporation,  leaving  the  salts  behind  and  forming 
the  highly  saline  areas  commonly  known  as  salt-spots.  In  the  salt-spots  the 
most  important  salts  are  those  of  sodium.  From  these  conditions  it  appears 
that  such  perennials  as  live  in  areas  where  the  salt  content  of  the  substratum 
is  relatively  high  must  not  only  be  able  to  extract  water  from  a  relatively 
highly  concentrated  soil  solution,  but  must  be  able  to  endure  the  salts  of  what- 
ever kind  as  such.  In  addition  to  these  conditions  the  salt  plants  live  in  a 
climate  otherwise  very  arid.  The  physiological  activities  of  halophytes  are 
accordingly  of  great  interest  in  that  they  appear  to  include  an  especially  high 
osmotic  efficiency  as  well  as  immunity  to  the  salts  of  sodium.  Their  diver- 
gence in  these  regards  from  many  desert  plants,  and  especially  from  meso- 
phytes,  is  thus  very  great. 

VEGETATION    OF   A    SALT-SPOT    NEAR   TUCSON. 

A  salt-spot  of  considerable  extent  lies  on  the  edge  of  the  flood-plain  of 
the  Santa  Cruz  river  along  the  old  Fort  Yuma  road  about  four  miles  north- 
west of  the  city  of  Tucson.  The  area  is  more  or  less  sharply  set  off  from  the 
surrounding  bottom  land  by  the  surface  depression,  but  especially  by  the 
high  salt  content  of  the  soil  and  by  the  halophytic  character  of  its  perennial 
plant  covering.  The  vegetation  of  the  non-salt  lands  adjoining  the  salt-spot 
is  made  largely  of  Prosopis  velutina  (the  mesquite),  two  or  three  Acacias, 
A  triplex  canescens,  Bigelovia  hartivegii,  Koerberlinia  spinosa,  Suceda  suffru- 
tescens  and  Zizyphus  lycioides.  Of  these  plants  Prosopis  and  Bigelovia  occur 
also  along  the  washes  which  traverse  the  salt-spot.  Among  the  plants  which 
are  typical  of  the  salt-spot  proper  are  at  least  four  species  of  A  triplex,  namely, 
canescens,  elegans,  nuttallii  and  polycarpa.  In  addition  to  these  there  may 
be  found  species  of  Lycium  and  small  specimens  of  Suceda  (Dondia). 

Generally  speaking,  the  salt  plants  have  a  well-marked  zonal  distribu- 
tion. As  one  enters  the  spot  he  encounters  A  triplex  canescens  and  no  other 
salt  bush.  But  associated  with  A.  canescens  are  small  specimens  of  Prosopis 
and  Bigelovia  in  abundance.  This  zone  is  referred  to  in  this  paper  as  the 
canescens  zone.  As  one  passes  through  this  zone  on  his  way  to  the  more  cen- 


124 


DUDLEY    MEMORIAL    VOLUME 


tral  portion  of  the  salt-spot  he  enters  the  polycarpa  zone,  where  such  non-salt 
loving  plants  as  are  found  there  are  along  the  washes  only.  Still  nearer  the 
center  of  the  spot  A.  nuttalli  is  met  and  with  it  A.  elegans  and  dwarfed 
specimens  of  Suceda  suffrutescens.  In  places  where  there  are  evidently  less 
salts,  species  of  Lycium  and  Prosopis  occur  along  the  washes.  The  inner 
area  is  called  here  the  nuttallii  zone.  At  the  very  center  of  the  spot  there  are 
no  plants  and  the  surface  of  the  ground  usually  shows  white  incrustations  of 
salts. 

CHARACTER  OF  THE  SOIL. 

The  soil  of  the  Santa  Cruz  river  bottoms  is  largely  "adobe,"  but  that  of 
the  salt-spot  is  a  fine  sandy  loam.  Mechanical  and  chemical  analyses  made 
by  the  Bureau  of  Soils,  U.  S.  Department  of  Agriculture,  of  samples  of  soil 
taken  from  the  center  of  the  spot  are,  for  the  upper  12  inches,  as  follows: 
Clay,  33.3%  ;  silt,  21.5% ;  very  fine  sand,  17.0%  ;  fine  sand,  23.5%  ;  medium 
sand,  2.6%;  coarse  sand,  2.0%;  fine  gravel  0.1%. 


Table  1.     Analysis  of  soil  from  salt-spot.1 


CONSTITUENTS 

CALCULATED    COMBINATIONS 

Ca 

trace 

MgS04 

3.38% 

Mg 

.70% 

Na2SO4 

70.78 

Na 

29.45 

KC1 

4.42 

K 

2.33 

NaCl 

5.94 

So4 

50.53 

NaHCO3 

15.48 

Cl 

5.70 

HC03 

11.29 

The  salts  are  not  uniformly  distributed  in  the  soil.  The  total  soluble 
salts  in  the  first  foot,  as  determined  by  resistance  tests  made  by  the  Bureau 
of  Soils,  is  1,72%;  in  the  second  foot,  1.0%,  and  in  the  third  foot,  1.3%. 
The  salt  content  was  not  observed  at  a  greater  depth  than  three  feet.  Ob- 
servations on  the  resistance  of  the  soil  solutions,  which  the  writer  made  with 
an  electric  bridge  of  the  type  used  by  the  Bureau  of  Soils2,  showed  that  the 


1  Analysis  furnished  by  the  Bureau  of  Soils,  U.  S.  Dept.  of  Agriculture.     The 
soil  was  from  1-12  inches  deep  taken  from  the  center  of  the  salt-spot,  Tucson. 

2  The    instrument    employed    in    these   tests    was    kindly   loaned   me   by    Prof. 
R.  H.   Forbes,  Arizona  Experiment   Station. 


SALT    PLANTS    AND    SALT-SPOTS CANNON 


125 


salts  are  likewise  unequally  distributed  horizontally.  Following  is  presented 
the  typical  resistance  of  soil  solutions  for  different  portions  of  the  salt-spot; 
the  results  have  not  been  reduced  to  percentages : 

Table  2.     Resistance  in  ohms  of  soil  solutions  of  salt-spot  near  Tucson. 


1ST    FOOT 

2o  FOOT 

3D    FOOT 

Canescens  zone 
Polycarpa  zone 
Nuttallii  zone 

1100   ohms 
160   ohms 
3203  ohms 

1350  ohms 
350  ohms 
31  ohms 

850  ohms 
225  ohms 
34  ohms 

The  field  tests  show,  then,  that  the  salts  are  most  abundant  in  the  nut- 
talli  zone,  and  least  abundant  in  the  canescens  zone,  and  in  the  zone  between 
the  two  they  are  intermediate  in  amount.  A  detailed  study  of  the  salt-spot 
would  show  a  secondary  variation  in  the  quantities  of  salts  in  the  soil,  as 
where  the  washes,  which  are  flooded  with  each  heavy  rain,  run  into  and 
through  the  spot.  In  such  places  the  soil  is  leached  and,  as  above  stated, 
makes  possible  the  invasion  of  less  resistant  salt  plants  and  even  of  non-salt 
plants. 

CHEMICAL  ANALYSES  OF  SALT  PLANTS. 

The  observed  zonal  distribution  of  the  salt  plants  taken  in  connection 
with  the  marked  vertical  and  horizontal  distribution  of  the  salts  in  the  soil, 
suggested  the  desirability  of  conducting  an  inquiry  into  the  character  of  the 
constituents  of  the  plants  themselves.  This  was  done  along  two  lines,  namely, 
by  chemical  analysis4,  and  electric  resistance. 


3  This  anomalous  condition  is  not  explained  in  the  notes.     In  dry  seasons  the 
ground  at  this  place  is  covered  with  an  incrustation  of  salts. 

4  The  chemical  analyses  were  made  by  Mr.  William  H.  Walker,  Laboratory  of 
Physiological  Chemistry,  Columbia  University,  and  through  the  kindness  of  Pro- 
fessor Wm.  J.   Gies. 


126 


DUDLEY    MEMORIAL   VOLUME 


Table  3.     Summary  of  analytic  results  in  percentages. 
(Analysis   of   William   H.   Walker.)5 


A  triplex  poly  car  pa 

Atriplex  nuttallii 

Atriplex  canescens 

CALCULATED 

CALCULATED 

CALCULATED 

FOUND     FOR  WATER 

FOUND     FOR  WATER 

FOUND     FOR  WATER 

FREE  ASH 

FREE  ASH 

FREE  ASH 

Si02 

1.93%       2.00% 

4.69%     4.72% 

1.68%     1.71% 

Fe2O3,  P2O5,  A12O3 

1  .  44         1  .  50 

2  .  63         2  .  65 

2.87         2.92 

Fe203 

0.31            .32 

0.44         0.44 

0.61         0.62 

P205 

1.10         1.15 

0  .  87         0  .  88 

1.89         1.92 

A1203 

0.03            .03 

1.32         1.33 

0.37         0.38 

CaO 

15.93       16.54 

8.27         8.33 

17.31       17.59 

MgO 

7  .  00         7  .  26 

3.66         3.69 

10.35       10.51 

S03 

3  .  70         3  .  84 

5.17         5.21 

8.40         8.56 

Cl 

9.11         9.43 

24.56       24.74 

9.15         9.30 

C02 

19.82       20.57 

11.67       11.76 

15.00       15.24 

Na2O 

18.88       19.60 

26.58       26.77 

1  .  74         1  .  80 

K2O 

18.56       19.26 

12.04       12.13 

31.85       32.37 

H2O  (by  difference) 

3.63 

0.73        

1  .  65        

The  following  supplemental  report  was  sent  later  by  Prof.  Gies  (April 
22,   1907)  : 


Table  4.     Results  obtained  by  washing  the  ashes  free  from  chloride. 


.      INSOLUBLE    MATTER 

SOLUBLE    MATTER 

Atriplex  nuttallii 
Atriplex  poly  car  pa 
Atriplex  canescens 

20.80% 
33.01% 
37.725% 

79.20% 
66.99% 
62.275% 

SUMMARY   OF   TESTS  OF   ELECTRIC   RESISTANCE   OF   PLANT   SOLUTIONS. 

The  electric  resistances  of  the  plant  solutions  were  determined  by  the  use 
of  the  same  apparatus  as  was  employed  in  testing  soil  solutions.  Several 
methods  were  used  in  preparing  the  material  for  testing,  as  follows:  Air  dry 
leaves  and  young  twigs  were  boiled  in  water  for  five  minutes,  and  the  solution 
expressed  by  the  use  of  a  small  press.  Fresh  leaves  and  young  twigs  were 


5  The  figures  in  the  first  of  each  pair  of  columns  are  averages  of  closely  agreeing 
results  in  duplicate.  The  second  column  of  each  pair  gives  the  figures  for  corre- 
sponding percentage  contents  in  water-free  material. 


SALT    PLANTS    AND    SALT-SPOTS CANNON 


127 


treated  in  a  similar  manner.  As  a  control,  a  portion  of  air-dry  material, 
which  had  been  boiled,  was  left  one-half  hour  in  cool  water,  after  which  the 
solution  was  expressed  as  usual.  The  method  usually  followed  was  to  bring 
the  plant  to  the  laboratory  and  allow  it  to  become  air-dry.  A  10%  solution 
was  then  prepared,  which  was  boiled  five  minutes.  The  fluid  was  expressed 
and  allowed  to  cool. 

Very  many  tests  were  made  mainly  on  the  salt  plants  named,  and  also 
on  other  non-salt  forms.  The  results  were  so  consistent  that  it  will  not  be 
necessary  to  give  more  than  a  single  representative  experiment. 


Table  5.     Electric  resistance  of  plant  solutions. 
Can  esc  ens  Zone. 


OHMS 

TEMPERATURE    OF 
SOLUTION 

Atriplex  canescens 
Bigelovia  hartwegii    ® 
Prosopis  velutina    ® 

65 
115 
400 

30°  0  C. 
28°  5  C. 
28°  0  C. 

Polycarpa   Zone. 

Atriplex  poly  car  pa 
Suceda  suffrutescens 
Bigelovia  hartwegii   ® 
Atriplex  canescens    ® 
Lycium 

35 
25 
110 
70 
67 

28.0°  C. 
id 
id 
id 
id 

Nuttallii  Zone. 

Atriplex  nuttallii 
Suceda  suffrutescens 
Prosopis  velutina    ® 

25 
42 
280 

28.0°  C. 

id 
id 

The  averages  of  five  resistance  tests  of  solutions  of  salt  plants,  made  at 
various  times  during  the  summer,  are  as  follows:  Atriplex  canescens,  41 
ohms;  Atriplex  polycarpa,  23.9  ohms;  Atriplex  nuttallii,  18.2  ohms7. 


6  These  plants  were  situated  in  some  of  the  small  washes  which  traversed  the 
salt-spot. 

7  A  verification  of  the  relative  results  obtained  by  the  use  of  the  bridge  was 
obtained  by  freezing  plant  solutions  previously  tested  electrically.     The  following 
were   some  of  the  results : 


128 


DUDLEY    MEMORIAL    VOLUME 


Atriplex  canescens 
Atriplex  poly  car  pa 
Atriplex  nuttallii 


ELECTRIC    RESISTANCE 

70.0  ohms 
29 . 0  ohms 
16.5  ohms 


FREEZING    POINT 

-0.2°  C. 
-0.3°  C. 
-0.4°  C. 


SOME   INFERENCES   AND   CONCLUSIONS,    AND   SUMMARY. 

The  most  important  findings  of  the  present  study,  together  with  sugges- 
tions as  to  their  possible  significance,  may  be  briefly  presented. 

Chemical  analyses  of  three  species  of  halophytes,  which  occur  naturally 
in  a  certain  salt-spot,  or  area,  where  there  is  a  large  amount  of  "white  alkali," 
near  Tucson,  show,  among  other  features,  that  the  amount  of  salts,  as  well 
as  the  kind,  is  unlike  and  is  characteristic  for  the  species.  It  is  found  also 
that  certain  elements,  especially  sodium  and  calcium,  are  present  in  unlike 
amounts  in  the  ash  of  these  plants.  The  relation  of  sodium  and  calcium  in 
the  salt  plants,  and  in  mesophytes,  based  on  data  already  presented,  is  given 
in  the  following  table: 


Table  6. 


Calcium  and  sodium  in  plants,  in  per  cent,  with  the 
Ca 


ratio 


Na" 


Ca 

Na 

Ca 

Na" 

Cultivated  plants  
Fresh  water  plants  

7.9 
43.75 
17  31 

3.1 
8.58 
1  74 

2.5 
5.1 
9  8 

15  93 

18  88 

84 

Atriplex  nuttallii                          ... 

8  27 

26  58 

31 

Saltwater  plants  

12.88 

24.81 

.31 

Electric  resistance  tests  were  carried  out  on  the  same  species  of  halo- 
phytes as  were  subjected  to  analysis.  These  showed  in  general  that  the  re- 
sistances were  characteristic  for  any  given  species.  Atriplex  canescens  had 
the  highest  and  A.  nuttallii  had  the  lowest  resistance. 

Physical  and  chemical  studies  of  the  alkali  soil  were  also  made.  Sodium 
salts  were  found  especially  abundant  and  calcium  salts  were  present  as  a  trace 
only.  The  salts  were  found  to  be  most  abundant  at  the  center  of  the  salt- 
spot  and  to  become  less  gradually  as  the  periphery  of  the  spot  was  ap- 
proached. 

Combining  the  observations  on  both  plants  and  soil  we  find,  therefore, 
that  the  center  of  the  salt-spot,  where  the  soil  solution  is  most  dense  and 


SALT    PLANTS    AND    SALT-SPOTS CANNON  129 

carries  the  most  sodium  salts,  is  inhabited  by  the  species,  A.  nuttallii,  which 
contains  the  largest  amount  of  soluble  salts  and  also  the  largest  proportion  of 
sodium.  It  has  also  the  least  amount  of  calcium.  The  species  which  occupy 
the  outer  portion  of  the  salt-spot,  where  the  salts  are  least  in  amount,  are 
those  which  contain  the  least  amount  of  sodium  and  the  largest  of  calcium. 
From  these  facts  it  is  inferred  that  there  is  probably  an  increase  in  the 
osmotic  pressure  in  the  different  species  as  one  goes  from  the  periphery  of  the 
salt-spot  to  the  center,  and  that  among  the  substances  which  contribute  to 
this  effect  the  salts  of  sodium  may  find  an  important  place.  Whether  there 
is  also  an  increase  in  such  osmotically  active  organic  substances  as  are  not 
electrolytes,  the  inference  necessarily  leaves  out  of  consideration. 

It  has  already  been  stated  that  the  relative  amounts  of  sodium  and  cal- 
cium in  the  different  species  of  salt  plants  are  unlike,  Na  being  most  abun- 
dant in  such  species  as  contain  the  lowest  proportion  of  Ca  and  vice  versa. 
These  facts  are  so  striking  that  they  may  have  a  special  bearing  beyond  the 
well  known  facility  of  plants  of  storing  up  in  insoluble  form  certain  salts. 
There  may  be  some  relation,  also,  with  the  occurrence  of  salts  in  the  soil 
solution.  It  is  known8  that  lime,  when  applied  to  certain  alkali  soils,  serves 
as  a  correction,  making  such  soils  more  tolerable  for  mesophytes.  It  should 
be  noted  that  in  such  conditions  the  soil  solution  is  probably  made  more  dense. 
Osterhout9  has  shown  that  calcium  holds  an  antagonistic  relation  to  sodium 
by  which  the  latter  may  be  prevented  from  entering  the  protoplast.  Applying 
these  findings  to  the  case  in  hand,  it  may  be  concluded  that  the  salt-spot  in 
question,  if  treated  with  a  proper  amount  of  calcium  salts,  might  support  an 
entirely  different  type  of  vegetation,  or  that  the  kinds  of  salt  plants  inhabit- 
ing it  would  be  different  than  at  present,  owing  probably  to  the  absence  of 
the  more  intensely  halophytic  species.  In  such  a  case,  the  density  of  the  soil 
solution  would  play  a  minor  role.  The  findings  of  the  present  study  also 
indicate  that  the  most  intense  halophytes  absorb  salts  of  sodium  in  large 
amounts  without  injury,  and  that  it  is  due  to  this  that  such  species  can  sur- 
vive where  such  salts  constitute  the  leading  features  of  the  substratum. 

8  Some  mutual   relations   between   alkali   soils   and   vegetation.    Kearney   and 
Cameron.    U.  S.  Dept.  Agric.  Rep.  No.  71,  1902. 

9  The  permeability  of  protoplasm  to  ions  and  the  theory  of  antagonism,  Science, 
N.  S.,  Vol.  35,  page  112,  1912.    The  permeability  of  living  cells  to  salts  in  pure 
and  balanced  solutions,  Science,  N.  S.,  Vol.  34,  page  187,  1911. 


NORTH  AMERICAN   SPECIES  OF  THE   GENUS  AMYGDALUS.1 

WILLIAM  FRANKLIN  WIGHT, 
Bureau  of  Plant  Industry,  Washington,  D.  C. 

THE  SPECIES  which  form  the  subject  of  the  present  paper  have  been 
referred  to  both  Prunus  and  Amygdalus  by  various  authors  who  have 
had  occasion  to  consider  them.  They  are,  however,  so  distinct  from 
all  other  species  of  Prunus  in  America  as  at  once  to  suggest  the  propriety 
of  constituting  a  new  genus.  This  in  fact  was  done  by  Torrey2  for  one 
species  of  the  group,  he  having  published  the  genus  Emplectocladus  in  1854. 
Notes  on  a  few  specimens  in  the  herbarium  of  the  Museum  d'Histoire  Nat- 
urelle  at  Paris  indicate  that  Spach  who  studied  the  Amygdalaceae  as  thor- 
oughly as  any  one  of  his  time,  also  had  under  consideration  the  recognition 
of  the  genus  Emplectocladus.  A  study  of  the  material  preserved  in  some 
of  the  European  herbaria,  however,  shows  conclusively  that  they  cannot  be 
separated  from  certain  Asiatic  species  which  are  referred  to  Amygdalus  by 
authors,  who  recognize  the  latter  as  a  genus  distinct  from  Prunus.  In  fact, 
the  Asiatic  species  apparently  show  a  gradual  but  complete  transition  from 
this  North  American  group  to  the  species  of  Amygdalus  common  in  cultiva- 
tion. Again,  while  the  groups  Padus  and  Laurocerasus  appear  to  be  sep- 
arable from  true  Prunus  by  well-defined  characters,  it  must  be  admitted  that 
when  all  the  species  are  considered,  the  groups  Prunus  and  Amygdalus  show 
no  such  well-marked  diagnostic  characters,  even  though  their  extremes  are 
different  enough.  Nevertheless,  unless  these  two  groups  are  recognized'  as 
separate  genera,  to  speak  or  write  of  a  given  species  as  belonging  to  Prunus 
conveys  little  conception  of  its  true  character,  or  to  what  economic  species  it 
may  be  most  closely  related.  Since  nomenclature  cannot  well  be  made  to 
express  accurately  the  relationship  of  species,  but  is,  first  of  all,  a  matter  of 
convenience,  the  present  author  would  retain  the  name  Amygdalus  for  the 
peach  and  almond-like  species.  And  while  this  group  may  not  be  readily 

1  In  the  preparation  of  this  paper  the  author  has  consulted  the  material  in  the 
following  herbaria:    the  Gray  Herbarium,  Arnold  Arboretum,  New  York  Botanical 
Garden,    United    States    National    Herbarium,    Missouri    Botanical    Garden,    Field 
Museum,  University  of  California,  Leland  Stanford  Jr.  University,  Kew  Gardens, 
and  the  Museum  d'Histoire  Naturelle  of  Paris.     The  curators  of  these  collections 
have  kindly  made  available  the  material  in  their  charge  and  for  this  the  writer 
wishes  to  express  his  appreciation. 

2  Torrey,  John,  Smithsonian  Contributions  VI.     (Plantae  Fremontianae  10.  t.  5. 
1854.) 


NORTH    AMERICAN    AMYGDALUS WIGHT  131 

separated  by  prominent  diagnostic  characters,  the  species  do  form  a  fairly 
natural  group  and  the  name  Amygdalus  at  once  conveys  to  the  mind  charac- 
ters very  different  from  those  associated  with  true  Prunus.  • 

AMYGDALUS  TEXANA  (Dietr.)  W.  F.  Wight. 
Amygdalus  glandulosa  Hook.  Ic.  PI.  3  :t.  288.   1840. 
Prunus  glandulosa  Torr.  &  Gray,  Fl.  N.  Am.  1 :  408.   1840. 
Prunus  texana  D.  Dietr.  Syn.  PI.  3:  45.   1843. 
Prunus  hookeri  Schneider,  Hanb.  Launholz.  1:  597.   1905. 

Leaves  oval  to  oblong-oval,  mostly  14  to  25  mm.  or  sometimes  30  mm. 
long  on  young  succulent  shoots,  6  to  15  mm.  broad,  slightly  narrowed  toward 
the  base,  mostly  obtuse  at  the  apex,  green  and  pubescent  above,  grayish  to 
mentose  below,  the  margins  conspicuously  glandular-serrate;  petioles  1  to  4 
mm.  long,  pubescent ;  stipules  linear  and  glandular.  Flowers  appearing  in 
March  with  or  slightly  before  the  leaves,  solitary  or  in  twos,  10  to  12  mm. 
broad ;  pedicels  3  to  4  mm.  long,  grayish  pubescent ;  calyx  pubescent,  the  tube 
about  2  mm.  long,  the  oblong-ovate  lobes  as  long  and  glandular-serrate; 
petals  apparently  white,  oblong,  4.5  to  6  mm.  long,  obtuse  at  the  apex  and 
abruptly  narrowed  to  a  very  short  claw.  Fruit  rather  densely  velvety-hairy, 
apparently  maturing  in  June;  stone  ovoid  13  to  15  mm.  long,  11  mm.  broad 
and  9  mm.  thick,  usually  rounded  at  the  base,  pointed  at  the  apex  and  slightly 
flattened  toward  the  ventral  edge  but  rounded  on  the  dorsal  side,  the  surface 
smooth. 

A  dwarf  bushy  shrub  with  very  irregular  branches  and  grayish  bark,  that 
of  the  young  branchlets  light-gray  and  strongly  pubescent.  It  occurs  only 
in  Texas,  where  it  is  found  in  granitic  soils  from  between  Laredo  and  Bejar, 
the  type  locality  of  Hooker's  material  preserved  in  the  Kew  Herbarium,  to 
Llano  and  Mason  Counties  and  eastward  to  Hampstead. 

The  specific  name  glandulosa  was  used  earlier  for  a  different  species  by 
Thunberg  and  the  name  hookeri  was  published  by  Schneider  for  this  reason. 
There  appears,  however,  to  be  nothing  to  prevent  the  use  of  Dietrich's  name 
texana,  which  was  doubtless  overlooked  by  Schneider.  The  original  descrip- 
tion by  Dietrich  was  based  on  a  specimen  collected  by  Drummond  but  for 
which  the  locality  is  not  given. 

Specimens  examined:  Texas ;  Stevens  Bend,  Colorado  River,  Lampasas 
Co.,  C.  S.  Mason,  1910.  Hoover's  Hill,  near  Kingston,  C.  S.  Mason,  Mar. 
26,  1910.  Llano  Co.,  Reverchon,  May,  1885.  Big  Branch,  Gillespie  Co., 
G.  Jermy.  Sandy  Plains,  Hampstead,  Elihu  Hall,  No.  189,  June  4,  1872, 
San  Antonio  Wells  on  the  Prairie  road  between  Ringgold  Banks  and  Laredo, 
G.  H.  Thomas,  Feb.,  1857.  Kimble  Co.,  I.  Reverchon,  No.  53,  May,  1885. 


132  DUDLEY    MEMORIAL    VOLUME 

Sandy  region  of  Llano  and  Mason  Co.,  I.  Reverchon,  May,  1885.     Between 
Laredo  and  Be  jar,  Berlandier. 

Without  locality,  Lindheimer,  No.  236,  1844,  Drummond.  Hall,  No. 
189,  1872.  Chas.  Wright,  1848. 

AMYGDALUS  MICROPHYLLA  H.  B.  &  K. 
Amygdalus  microphylla  H.  B.  &  K.  Nov.  Gen.  &  Sp.   PI.  6:  243.  t.  564. 

1823. 
Prunus  microphylla  Hemsl.  Biol.  Centr.  Am.  Bot.  1:  368.   1879-88. 

Leaves  with  petioles  about  1  mm.  long,  oblong-oval  to  oblong-oblanceo- 
late,  10  to  15  mm.  long,  3  to  6  mm.  broad,  glabrous  on  both  surfaces  at  matur- 
ity but  when  immature  often  sparingly  pubescent,  narrowed  toward  the  base, 
obtuse  at  the  apex,  the  margins  crenate-dentate  and  glandular  when  young, 
the  glands  in  age  remaining  as  callous  mucronate-like  points.  Flowers  ap- 
pearing with  or  before  the  leaves  on  very  short  lateral  spurs,  one  to  three  on 
each  spur,  the  pedicels  pubescent  and  1  mm.  or  less  long;  calyx  turbinate, 
about  3  mm.  long,  glabrous,  the  lobes  1  mm.  long,  obtuse  and  furnished  with 
a  few  marginal  glands;  petals  oval  or  obovate,  obtuse,  1.5  mm.  long  and  1 
mm.  broad.  Fruit  in  dried  specimens  about  12  mm.  long,  9  mm.  in  diameter, 
oblong-ovoid,  rounded  at  the  base,  terminated  at  the  apex  by  a  slight  mucro- 
nate-like point;  the  exocarp  dry  and  pubescent,  dehiscent  along  the  ventral 
side;  stone  about  10  mm.  long,  8mm.  broad,  slightly  winged  on  the  ventral 
side  and  with  an  indistinct  groove  along  the  dorsal  edge,  the  surface  smooth. 

A  small  shrub  with  rather  rigid  branches  and  more  or  less  spinescent 
branchlets,  the  bark  of  the  young  growth  light  gray  and  pubescent  but  be- 
coming dark  gray  with  age.  The  species  was  originally  described  from 
Mexico  on  dry  hills  between  Pachuca  and  Moran  at  7,800  feet  altitude.  It 
seems  to  have  been  rarely  collected,  as  very  little  material  is  to  be  found  in 
herbaria,  either  in  Europe  or  America. 

Specimens  examined:  Mexico;  chiefly  in  the  region  of  San  Luis  Potosi, 
6,000  to  8,000  ft.  alt.,  C.  C.  Parry  and  E.  Palmer,  No.  221,  1878.  Prope 
El  Gigante,  Hartweg,  No.  1602. 

AMYGDALUS  MINUTIFLORA  (Engelm.)   W.  F.  Wight. 

Prunus  minutiflora  Engelm.;  Gray,  Bost.  Journ.  Nat.  Hist.  6:  185.    1850. 
Cerasus  minutiflora  Gray,  PL  Wright,  pt.  1.  68.   1852. 

Leaves  oblong-oval,  narrowed  toward  the  2  to  3  mm.  long  petiole  and 
rounded  at  the  apex,  7  to  20  mm.  long,  5  to  8  mm.  broad,  pale  below  and 
glabrous  on  both  surfaces,  the  margins  sometimes  slightly  revolute,  entire  or 
occasionally  irregularly  toothed,  glandless.  Flowers  appearing  with  the 
leaves  in  February  or  March  on  short  lateral  spurs,  sessile  or  very  nearly  so, 


NORTH    AMERICAN   AMYGDALUS  —  WIGHT  133 

one  to  four  on  each  spur;  calyx-tube  campanulate  with  broadly  ovate  lobes 
about  1  mm.  long;  petals  white,  obovate,  about  3.5  mm.  long.  Fruit  matur- 
ing in  June,  globose,  about  12  mm.  long,  11  mm.  broad  and  10  mm.  thick 
when  dry,  the  exocarp  thin  and  dehiscing  along  the  ventral  edge ;  stone  turgid 
and  with  a  smooth  surface,  grooved  along  the  dorsal  edge,  the  ventral  edge 
rather  thick  with  an  inconspicuous  groove  a  short  distance  from  the  margin. 

A  low  shrub  with  irregular,  often  zigzag,  scarcely  spinescent  branches, 
and  grayish  bark.  It  sometimes  forms  dense  masses  and  occurs  in  Texas 
from  the  vicinity  of  New  Braunfels  westward  to  Devils  River  and  the  Rio 
Grande.  It  was  originally  described  from  "Hills  and  dry  slopes  between 
San  Antonio  and  New  Braunfels." 

Specimens  examined:  Texas;  Comanche  Spring,  New  Braunfels,  Lind- 
heimer,  No.  789,  Mar.,  1850.  San  Antonio,  Mackensen,  Feb.  20,  and  June, 
1910.  Valvada  Co.,  C.  S.  Mason,  May  10,  1910.  Between  New  Braunfels 
and  San  Antonio,  Lindheimer,  No.  401,  Mar.,  1850.  San  Antonio,  Geo. 
Thurber,  Mar.,  1853.  San  Felipe  Creek,  C.  Wright,  1851.  Without  local- 
ity, Lindheimer,  No.  388. 

AMYGDALUS  HARVARDII,  W.  F.  Wight,  sp.  nov. 

Leaves  obovate  to  oblong-obovate  or  sometimes  fan-shaped  on  young 
growth,  7  to  20  mm.  long,  3  to  10  mm.  broad,  glabrous  or  sometimes  finely 
pubescent  on  both  surfaces,  usually  somewhat  pale  below  and  under  a  lens 
rather  prominently  reticulate  veined,  the  margin  conspicuously  dentate  toward 
the  apex,  very  rarely  toothed  below  the  middle,  the  teeth  usually  acute  and 
apparently  glandless.  Flowers  appearing  with  the  leaves  and  sessile;  calyx 
slightly  pubescent,  the  tube  about  2.5  mm.  long,  the  lobes  scarcely  more 
than  1  mm.  long,  entire  and  obtuse;  petals  not  seen.  Fruit  sessile,  nearly 
globular,  the  pubescent  exocarp  dehiscent  along  one  edge,  when  dry  about  9 
mm.  long,  7  mm.  broad  and  7.5  mm.  thick;  stone  about  8  mm.  long,  6.5  mm. 
broad  and  7  mm.  thick,  rounded  at  the  base  and  slightly  pointed  toward  the 
apex,  the  surface  smooth  except  for  indistinct  grooves  near  the  ventral  edge. 

A  shrub  with  rather  rigid  branches,  stout  spinescent  branchlets  and  light 
gray  bark.  The  type  specimen  in  the  United  States  National  Herbarium 
was  collected  in  fruit  by  V.  Havard  in  July,  1883,  in  western  Texas,  east  of 
the  Chisas  Mountains,  near  Bone  Springs.  It  was  also  collected  by  C.  C. 
Parry,  J.  M.  Bigelow,  Charles  Wright  and  A.  Schott  on  the  Mexican  Boun- 
dary Survey  under  the  direction  of  Major  W.  H.  Emery,  this  specimen  being 
labeled  "chiefly  in  the  valley  of  the  Rio  Grande,  below  Donana."  The 
species  is  most  closely  related  to  Amygdalus  microphylla  H.  B.  &  K.  of 
Mexico,  but  is  easily  distinguished  by  its  broader,  more  obovate  leaves  as 
well  as  by  their  reticulate  venation  and  eglandular  margins. 


134  DUDLEY    MEMORIAL   VOLUME 

AMYGDALUS  FASCICULATA   (Torr.)    Greene. 

Amygdalus  fasciculata  Greene,   Fl.  Franciscana  49.     1891. 
Emplectocladus  fasciculata  Ton.  Smith.  Contr.  VI,  (PI.  Fremont.     10.  t.  5. 

1854.) 
Prunus  fasciculata  Gray,  Proc.  Am.  Acad.  10:  70.     1874. 

Leaves  oblanceolate-spatulate,  6  to  12  mm.  long,  1.5  to  2  mm.  broad, 
gradually  narrowed  toward  the  base  to  an  indistinct  petiole,  obtuse  or  some- 
times acute  at  the  apex,  rather  pale  green  and  hirsute  pubescent  on  both 
surfaces  or  sometimes  very  sparingly  pubescent,  only  the  midvein  apparent, 
the  margin  entire  or  very  rarely  somewhat  dentate  and  the  teeth  either 
glandular  or  eglandular.  Flowers  about  6  mm.  in  diameter,  1  to  3  on  short 
spurs  and  sessile;  the  calyx  glabrous  without  but  hairy  within,  the  tube 
about  2  mm.  long,  the  lobes  1.5  mm.  long;  petals  apparently  white,  oblan- 
ceolate.  Fruit  ovoid,  when  dry  8  to  10  mm.  long,  7.5  mm.  broad  and  7  mm. 
thick  borne  on  a  pedicel  1.5  mm.  long,  the  dry  exocarp  pubescent  and 
dehiscent  along  one  edge;  stone  about  8.5  mm.  long,  6.5  mm.  broad  and 
6  mm.  thick,  rounded  at  the  base,  pointed  at  the  apex,  somewhat  winged  and 
with  a  shallow  groove  on  the  ventral  edge,  the  surface  smooth. 

A  shrub  with  grayish  or  often  whitish  bark,  numerous  straight,  scarcely 
spinescent  branches,  and  sometimes  pubescent  young  growth.  The  specimens 
from  which  this  species  was  originally  described  were  collected  by  Colonel 
Fremont,  but  their  labels  had  been  lost  and  the  locality  from  which  they 
were  obtained  is  therefore  unknown,  though  the  author  of  the  species  says 
"Sierra  Nevada  of  California;  probably  in  the  southern  part  of  the  range." 
The  species  is  known  to  occur  from  southern  Utah  through  southern  Nevada 
and  northern  Arizona  to  the  slopes  of  the  southern  Sierra  Nevada,  and  the 
San  Bernardino  Mountains  of  California.  A  specimen  has  also  been  seen 
labeled  "Santa  Maria,  Santa  Barbara  County,  California." 

Specimens  examined:  Utah;  southern  Utah,  C.  C.  Parry,  1874.  St. 
George,  at  2000  ft.  alt.,  M.  E.  Jones  No.  1630,  April  5,  1880.  Southern 
Utah,  Dr.  Palmer  No.  135,  1877.  Valley  of  the  Virgin,  C.  C.  Parry  No.  56, 
1874.  St.  George,  Dr.  Palmer,  1871.  Silver  Reef,  M.  E.  Jones,  1894. 

Nevada;  Morman  Mts.,  Lincoln  Co.,  at  6000  ft.  alt.,  Kennedy  and 
Gooding  No.  140,  July,  1906.  Rocky  hillsides,  Calientes,  L.  N.  Gooding 
No.  609,  April  26,  1902.  Kernon,  L.  N.  Gooding  No.  652,  April  28,  1902. 
Washes  in  the  Palmetto  Range  at  6-7000  ft.  alt.,  C.  A.  Purpus  No.  5852, 
May-Oct.,  1898. 

Arizona;  On  trip  to  Castle  Rock,  near  Tucson,  D.  Griffiths  No.  2119, 
Nov.  17,  1900.  Without  locality,  Dr.  Palmer,  1876.  Choride,  at  4500  ft. 
alt,  M.  E.  Jones,  April  14,  1903. 


NORTH    AMERICAN    AMYGDALUS WIGHT  135 

New  Mexico;  "Western  New  Mexico,"  Dr.  Bigelow  on  the  Whipple 
Expedition. 

California;  Panamint  Mts.,  Inyo  Co.,  Coville  &  Funston  No.  555,  April 
4,  1891.  Mill  Creek  Canyon,  Panamint  Mts.,  at  about  4200  ft.  alt.,  Coville 
&  Funston  No.  801,  May  20,  1891.  Lone  Pine  Canyon,  desert  slopes  of  the 
San  Gabriel  Mts.  at  4500  ft.  alt.,  Abrams  and  MacGregor  No.  668,  July  5, 
1908.  Morongo  Mts.,  Colorado  Desert,  S.  B.  &  W.  F.  Parish,  April,  1882. 
Mountains,  Mojave  Desert,  S.  B.  &  W.  F.  Parish,  May,  1882.  Sandy  places, 
south  fork  of  Kern  River,  at  2-3000  ft.  alt.,  C.  A.  Purpus  No.  5022,  1897. 
Santa  Maria,  Santa  Barbara  Co.,  L.  Jared,  1882.  Summit  of  Providence 
Mts.,  at  5000  ft.  alt.,  J.  G.  Cooper,  May  29,  1861.  San  Bernardino  Mts., 
Pipe  Canyon,  S.  B.  Parish,  June  16,  1894.  Pleasant  Canyon,  Panamint  Mts., 
5500  ft.  alt.,  M.  E.  Jones,  May  6,  1897.  Summit  Cajon  Pass,  San  Ber- 
nardino Co.,  at  4200  ft.  alt.,  H.  M.  Hall  and  H.  P.  Chandler  No.  6755, 
April  27,  1906.  San  Gabriel  Mts.,  Los  Angeles  Co.,  L.  R.  Abrams  and 
E.  A.  McGregor  No.  525,  July  3,  1908. 

AMYGDALUS  ANDERSONII   (Gray)   W.  F.  Wight. 
Prunus  andersonii  Gray,  Proc.  Am.  Acad.  7 :  337.     1868. 

Leaves  sessile,  oblong-oval  to  lanceolate,  sometimes  varying  to  oblan- 
ceolate  on  flowering  shoots,  10  to  20  mm.  long,  mostly  3  to  6  mm.  broad, 
cuneate  toward  the  base  and  furnished  with  one  to  three  glands  or  egland- 
ular,  acute  at  the  apex  or  sometimes  obtuse,  light  green  and  glabrous  on  both 
surfaces  or  slightly  paler  below,  the  margins  with  indistinct  acute  and 
eglandular  serrations  or  entire.  Flowers  appearing  in  March  and  April, 
about  18  mm.  broad  in  dried  specimens,  apparently  solitary  in  pedicels 
3  to  4  mm.  long;  the  calyx  glabrous,  the  tube  broadly  campanulate,  about 
2  mm.  long,  the  lobes  1.5  mm.  long,  obtuse,  pubescent  within,  the  margins 
sparingly  glandular  or  eglandular;  petals  pinkish,  obovate  or  oblong  and 
cuneate  toward  the  base,  6  to  8  mm.  long,  3  to  4  mm.  broad;  ovary  hirsute. 
Fruiting  pedicels  rather  stout,  6  to  10  mm.  long;  the  fruit  with  a  dry  exocarp 
and  pubescent  but  not  densely  so,  slightly  flattened  and  narrowed  toward  the 
base,  usually  rounded  at  the  apex  but  furnished  with  a  mucronate  point, 
when  dry  12  to  17  mm.  long,  10  to  14mm.  broad;  stone  11  to  13  mm.  long, 
9  to  11  mm.  broad  and  6  to  7  mm.  thick,  narrowed  toward  the  base,  obtuse 
at  the  apex,  within  a  rather  well-marked  ventral  wing  and  a  shallow  groove 
along  the  dorsal  edge,  the  surface  reticulate. 

A  shrub  about  1  m.  high  with  rather  rigid,  strict  branches  arid  more  or 
less  spinescent  branchlets,  the  young  growth  greenish  or  sometimes  with  a 


136  DUDLEY    MEMORIAL    VOLUME 

glaucous  appearance  turning  reddish  gray  with  age.  Originally  described 
from  the  "Foothills  of  the  eastern  side  of  the  Sierra  Nevada,  near  Carson." 
It  appears  to  be  rather  common  in  the  eastern  Sierra  Nevada  and  the  foot- 
hills of  both  California  and  Nevada  from  the  vicinity  of  Reno  in  Nevada 
southward  to  Owens  Valley  in  California. 

Specimens  examined:  Nevada;  Hawthorne,  at  6000  ft.  alt.,  M.  E. 
Jones,  April  15,  1907.  Verdi,  Washoe  Co.,  C.  F.  Sonne,  May,  1895.  Foot- 
hills northwest  of  the  University  Reno,  P.  B.  Kennedy,  April  22,  1901. 
Near  Carson  City,  C.  L.  Anderson,  1866.  Near  Empire  City,  J.  Torrey  No. 
136,  1865.  Carson  City,  at  5000  ft.  alt.,  S.  Watson  No.  299,  April,  1868. 
King's  Canyon,  Ormsby  Co.,  Nev.,  at  about  6000  ft.  alt.,  C.  F.  Baker  No. 
907,  June  1  and  July  1,  1902.  Empire  City,  Nevada,  M.  E.  Jones  No. 
3856,  May  19,  1882.  Candelaria,  W.  H.  Shockley,  1890.  Carson  Valley, 
J.  C.  Phillips  and  C.  S.  Sargent,  Sept.-Oct.,  1878.  Reno,  F.  H.  Hillman, 
Oct.,  1893.  Miller  Mts.,  at  7500  ft.  alt.,  W.  H.  Shockley  No.  1216,  May 
18,  1882.  Near  Carson  City,  C.  L.  Anderson  No.  217,  1864  and  Mar.,  1865. 
Near  Woodford,  border  of  Nevada,  J.  Ball,  July,  1884.  Carson  Flats,  J.  D. 
Hooker  and  Asa  Gray,  1877.  Reno,  J.  D.  Hooker  and  Asa  Gray,  1877. 

California;  White  Mts.,  at  5-6000  ft.  alt.,  C.  A.  Purpus  No.  5805, 
May-Oct,  1898.  Lone  Pine,  at  7000  ft.  alt.,  M.  E.  Jones,  May  14,  1897. 
Base  of  White  Mts.  east  of  Laws,  Inyo  Co.,  A.  A.  Heller  No.  8186,  May  5, 
1906.  Beckwith  Pass,  H.  N.  Bolander.  Camp  Bidwell,  D.  W.  Mathews, 
1879.  Owens  Valley  desert,  Dr.  W.  Matthews,  April,  1877. 

AMYGDALUS  FREMONTI  (S.  Wats.)   Abrams. 

Amygdalus  fremonti  Abrams,  Bull.  N.  Y.  Bot.  Card.  6:  384.     1911. 
Prunus  fremonti  S.  Wats.  Bot.  Calif.  2 :  442.     1880. 

Leaves  with  petioles  3  to  4  mm.  long,  ovate  to  nearly  orbicular,  rounded 
or  sometimes  subcordate  at  the  base,  obtuse  or  acute  at  the  apex,  12  to 
25  mm.  long,  10  to  18  mm.  broad,  pale  green  above,  grayish  or  sometimes 
even  somewhat  silvery  below  and  marked  with  reddish  brown  veins,  glabrous 
on  both  surfaces,  the  margins  strongly  crenate-dentate  and  the  teeth  gland- 
ular. Flowers  appearing  in  March  with  the  leaves,  solitary  or  in  fascicles 
of  two  or  more  on  pedicels  8  to  12  mm.  long;  calyx  with  ciliate  lobes;  the 
corolla  about  12  or  15  mm.  broad;  the  ovary  densely  pubescent.  Fruit 
pedunculate,  about  12  mm.  in  diameter,  the  exocarp  apparently  dry  and  de- 
hiscing along  the  ventral  side ;  stone  oblong  and  turgid,  about  1 2  mm.  long, 
10  mm.  broad,  and  8  mm.  thick,  rounded  on  the  dorsal  side  and  with  a 
thick  wing  on  the  ventral  side. 


NORTH    AMERICAN    AMYGDALUS WIGHT  137 

A  shrub  or  small  tree  5  m.  high  with  irregular  more  or  less  spinescent 
branches  and  reddish  bark,  originally  described  from  "Coast  Ranges  of 
Southern  California;  Oriflamme  Canon,  San  Diego  County  (D.  Cleveland)  ; 
San  Bernardino,  Parry  and  Lemmon,  n.  108,  1876.  Also  collected  by  Fre- 
mont in  1846,  locality  uncertain."  The  species  occurs  in  southern  Cali- 
fornia from  the  Chuckawalla  Mountains  southward  to  northern  Lower  Cali- 
fornia. 

The  type  sheet  of  this  species  in  the  Gray  Herbarium  at  Cambridge 
contains  with  other  specimens  a  fragmentary  one  collected  by  Fremont 
probably  in  1846,  but  this  is  Primus  subcordata  and  was  evidently  taken  by 
Watson  to  be  identical  with  the  other  material  and  Fremont's  name  was  ac- 
cordingly given  to  the  species,  though  it  is  doubtful  if  he  ever  collected  it. 

Specimens  examined:  California;  Palm  Springs,  C.  S.  Mason,  Mar., 
1908.  Agua  Caliente  Canyon,  C.  C.  Parry,  April,  1881.  Mountains  on  the 
edge  of  the  desert,  San  Diego  County,  Parish,  Mar.,  1881.  San  Felipe, 
C.  C.  Parry,  June,  1850.  Colorado  Desert,  W.  G.  Wright  No.  198,  Mar., 
1881.  Oriflamme  Canyon,  San  Diego  Co.,  D.  Cleveland,  1877.  Without 
locality,  Dr.  A.  Davidson,  1893.  Parry  and  Lemmon  No.  108,  1876. 

Lower  California;  Northern  Lower  California,  C.  R.  Orcutt,  April  12, 
1886.  Santo  Thomas,  C.  R.  Orcutt,  Sept.  29,  1884.  El  Rancho  Vego, 
T.  S.  Brandegee,  April  29,  1889. 


DATE  DUE 


Bio-Ag   QK3   D8    1913 
Dudley   memorial    volume. 


000657018     8 


